孕穗期低温对寒地不同水稻材料光合系统的影响研究

doi:10.16819/j.1001-7216.2025.241014

中国水稻科学 ›› 2025, Vol. 39 ›› Issue (5): 679-689.DOI: 10.16819/j.1001-7216.2025.241014

孕穗期低温对寒地不同水稻材料光合系统的影响研究

丁国华¹^,³^,^#, 李鑫²^,^#, 曹良子¹, 周劲松¹, 雷蕾¹, 白良明¹, 洛育¹, 杨光¹, 崔志波³, 赵明辉³^,^*(), 孙世臣¹^,^*()

¹黑龙江省农业科学院耕作栽培研究所/黑龙江省水稻品质改良与遗传育种工程技术研究中心/黑龙江省寒地作物生理生态重点实验室，哈尔滨 150086
²东北农业大学农学院，哈尔滨 150030
³沈阳农业大学东北粳稻遗传改良与优质高效生产省部共建协同创新中心，沈阳 110866

收稿日期:2024-10-31 修回日期:2025-03-25 出版日期:2025-09-10 发布日期:2025-09-10
通讯作者: *email: mhzhao@syau.edu.cn;email: sunshichen1979@163.com
作者简介:^#共同第一作者
基金资助:
黑龙江省省属科研业务费项目(CZKYF2023-1-C012);沈阳农业大学东北粳稻遗传改良与优质高效生产省部共建协同创新中心开放课题）(KF2022-04);国家水稻产业技术体系专项(CARS-01-62);黑龙江省农业科学院创新工程资助项目(CX23ZD02)

Effect of Low Temperature at Booting Stage on Photosynthetic System of Different Rice Materials in Cold Region

DING Guohua¹^,³^,^#, LI Xin²^,^#, CAO Liangzi¹, ZHOU Jinsong¹, LEI Lei¹, BAI Liangming¹, LUO Yu¹, YANG Guang¹, CUI Zhibo³, ZHAO Minghui³^,^*(), SUN Shichen¹^,^*()

¹Institute of Crop Cultivation and Tillage, Heilongjiang Academy of Agricultural Sciences/Heilongjiang Engineering Technology Research Center of Rice Quality Improvement and Genetic Breeding/ Heilongjiang Provincial Key Laboratory of Crop Physiology and Ecology in Cold Region, Harbin 150086, China
²College of Agriculture, Northeast Agricultural University, Harbin 150030, China
³Collaborative Innovation Center for Genetic Improvement and High Quality and Efficiency Production of Northeast Japonica Rice in China, Shenyang Agricultural University, Shenyang 110866, China

Received:2024-10-31 Revised:2025-03-25 Online:2025-09-10 Published:2025-09-10
Contact: *email: mhzhao@syau.edu.cn;email: sunshichen1979@163.com

摘要/Abstract

摘要： 【目的】解析龙稻18(LD18)和龙稻17029(L9)孕穗期光合系统对低温反应差异，为培育孕穗期耐冷高产水稻品种提供理论依据。【方法】以优质耐冷品种LD18和其与旱稻创制的高产冷敏感新种质L9为试验材料，利用人工气候室研究了孕穗期低温冷害对不同耐冷材料剑叶叶绿体超微结构、光合参数、相关酶活性及基因表达的影响。【结果】测序分析表明，LD18和L9基因组相似程度为75.03%，大的差异片段主要在2号、5号染色体。孕穗期低温下，LD18空壳率显著低于L9，LD18剑叶叶色值和叶绿体超微结构变化不明显，L9叶色值显著降低，叶绿体和类囊体出现膨大、变形，并产生了大量嗜锇粒，LD18光合相关酶活性高于L9。转录组及RT-qPCR结果表明，LD18差异表达基因数多于L9；GO和KEGG分析显示，LD18和L9相比最富集的基因通路包括光合作用和叶绿素代谢； RT-qPCR分析结果表明光合相关基因RBCX1在LD18中上调表达3倍，而在L9中下调表达。【结论】孕穗期低温下，LD18比L9具有更强的耐冷性，LD18能够调动更多基因应对低温胁迫，保持叶绿体结构完整，不降低叶绿素含量及光合速率，光合相关酶活性更高。

关键词: 冷害, 孕穗期, 生理性状, 光合作用, 转录组

Abstract:

【Objective】The study aims to analyze the reasons for differences in photosynthetic system response to low temperature of early maturing rice materials at booting stage in cold regions, and to provide theoretical and material basis for breeding cold-tolerant and high-yield rice varieties at booting stage.【Method】Using high-quality cold-tolerant variety Longdao 18 (LD18) and the new cold-sensitive high-yielding germplasm Longdao 17029 (L9) derived from crosses between LD18 and upland rice as experimental materials, we studied the effects of chilling injury on flag leaf chloroplast ultrastructure, photosynthetic parameters, related enzyme activities, and gene expression in materials with different cold tolerance.【Result】Genomic analysis showed 75.03% similarity between LD18 and L9, with major differences on chromosomes 2 and 5. Under low temperature during booting-stage, LD18 had a significantly lower unfilled grain rate than L9. The SPAD value and chloroplast ultrastructure of LD18 flag leaves showed no significant changes, while L9 exhibited significantly decreased SPAD value, swollen and distorted chloroplasts and thylakoids, and produced abundant osmiophilic granules. Photosynthetic-related enzyme activities were higher in LD18 than those in L9. Transcriptomic and RT-qPCR results indicated more differentially expressed genes in LD18 than in L9. GO and KEGG analyses revealed that the most enriched pathways in LD18 versus L9 included photosynthesis and chlorophyll metabolism. RT-qPCR showed 3-fold upregulation of the photosynthesis-related gene RBCX1 in LD18 but downregulation in L9.【Conclusion】At booting stage under low temperature, LD18 exhibits stronger cold tolerance than L9. LD18 can mobilize more genes to cope with low-temperature stress, maintain chloroplast structural integrity, avoid reduction in chlorophyll content and photosynthetic rate, and maintain higher photosynthetic enzyme activities.

Key words: chilling injury, booting stage, physiological trait, photosynthesis, transcriptome

丁国华, 李鑫, 曹良子, 周劲松, 雷蕾, 白良明, 洛育, 杨光, 崔志波, 赵明辉, 孙世臣. 孕穗期低温对寒地不同水稻材料光合系统的影响研究[J]. 中国水稻科学, 2025, 39(5): 679-689.

DING Guohua, LI Xin, CAO Liangzi, ZHOU Jinsong, LEI Lei, BAI Liangming, LUO Yu, YANG Guang, CUI Zhibo, ZHAO Minghui, SUN Shichen. Effect of Low Temperature at Booting Stage on Photosynthetic System of Different Rice Materials in Cold Region[J]. Chinese Journal OF Rice Science, 2025, 39(5): 679-689.

图/表 12

表1 RT-qPCR所用引物

Table 1. Primers used for RT-qPCR in this study

基因ID Gene ID	序列 Sequence(5' to 3')
18S-F	CTACGTCCCTGCCCTTTGTACA
18S-F	ACACTTCACCGGACCATTCAA
LOC4327351-F	ATTAGCCATCAACGACCA
LOC4327351-R	ACTCTCCAATCTGCTCTTC
LOC4331490-F	TGGCACAACTTACCTCTG
LOC4331490-R	CGCACATCTTCTACATCCT
LOC4333690-F	AACTGGAGGAACTGATTGG
LOC4333690-R	CACTGGACACGATAACGA
LOC4335640-F	GTCCCTTTACGACAATGGT
LOC4335640-R	CACCCGTGCTATCCATTAG
LOC4345611-F	GAGGTCCGATCTGCTTATG
LOC4345611-R	GCTGGTCTCCAAGACATAG
LOC4352460-F	ATGGCTTGTAGAGTAATGCT
LOC4352460-R	CTCTGCTGATCTGAACTCAT

图1 LD18（A）、L9（B）纯度及基因组相似度（C）

Fig. 1. LD18 (A), L9 (B) purity and genomic similarity (C)

图2 孕穗期低温下LD18和L9剑叶叶绿体超微结构（A）、叶色值（B）及空壳率（C） ns：无显著差异； *：P<0.05；***：P<0.001。T表示低温处理(17℃)；CK表示常温处理(25℃)。下同。

Fig. 2. Changes in chloroplast ultrastructure, SPAD value of flag leaves and percentage of unfilled grains for LD18 and L9 at low temperature during booting stage ns, No significant difference; *, P<0.05; ***, P<0.001.T, Chilling treatment((17℃), CK, Normal temperature treatment(25℃).

图3 孕穗期低温下LD18和L9剑叶光合速率及碳同化、能量代谢相关酶活性 ns：无显著差异； *：P<0.05；**：P<0.01。

Fig. 3. Activities of enzymes involved in carbon assimilation and energy metabolism in LD18 and L9 exposed to cold stress at the booting stage ns, No significant difference; *, P<0.05; **, P<0.01.

表2 转录组测序数据统计

Table 2. Transcriptome sequencing data statistics

样品名称 Sample	GC含量 GC content (%)	≥Q30百分比 ≥Q30 percent (%)	比对片段 Comparison fragment (bp)	比对到参考基因组 Compared to the percentage of the reference genome(%)
LD18-T1	52.39	94.46	41,811,440	95.93
LD18-T2	52.81	94.30	42,371,082	95.66
LD18-T3	52.59	94.42	41,630,423	95.60
LD18-CK1	53.16	94.50	37,394,980	95.58
LD18-CK2	53.06	94.53	43,242,371	95.01
LD18-CK3	52.63	94.93	39,951,714	96.15
L9-T1	52.76	94.89	37,865,138	95.43
L9-T2	51.66	94.19	44,570,598	95.03
L9-T3	52.62	92.88	41,324,819	94.96
L9-CK1	53.03	93.87	36,590,085	94.88
L9-CK2	52.84	93.92	40,478,494	95.25
L9-CK3	52.52	93.29	41,705,157	94.54

表3 孕穗期低温下两个材料发生差异表达基因数量

Table 3. Number of differentially expressed genes of the two materials at booting stage under cold stress

样品 Sample	差异表达基因数量 Number of DEGs	上调基因数量 Up-regulated genes	下调基因数量 Down-regulated genes
LD18-CK vs LD18-T	4509	2243	2266
L9-CK vs L9-T	3093	1432	1661

图4 孕穗期低温处理后LD18和L9均上调（A）和下调（B）表达的基因数量

Fig. 4. Up-regulated (A) and down-regulated (B) genes in LD18 and L9 compared with control at booting stage under low temperature

图5 孕穗期低温处理后LD18（A）和L9（B）中差异表达基因的GO分析

Fig. 5. GO analysis on DEGs in LD18 and L9 under cold stress relative to CK A：LD18-CK vs LD18-T；B：L9-CK vs L9-T.

图6 孕穗期低温处理后LD18（A）和L9（B）中差异表达基因的GO功能网络

Fig. 6. GO functional network map of DEGs in LD18 and L9 under cold stress relative to CK A：LD18-CK vs LD18-T；B：L9-CK vs L9-T.

图7 孕穗期低温处理后LD18（A）和L9（B）中差异表达基因的KEGG分析

Fig. 7. KEGG analysis on DEGs in LD18 and L9 under cold stress relative to CK A：LD18-CK vs LD18-T；B：L9-CK vs L9-T.

图8 LD18、L9中25个光合相关基因（A）和L9中6个光合相关差异表达基因（B）热图每个框中数字为归一化的表达量。

Fig. 8. Heat maps of 25 photosynthesis-related genes in LD18 and L9 (A) and 6 differentially expressed photosynthesis-related genes in L9 (B) Numbers in each box are normalized expressions.

图9 孕穗期低温下，LD18和L9中部分差异基因表达分析 ns：无显著差异； *：P<0.05；**：P<0.01。

Fig. 9. RT-qPCR validation of DEGs in LD18 and L9 due to CS ns, No significant difference; *, P<0.05; **, P<0.01.

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孕穗期低温对寒地不同水稻材料光合系统的影响研究

Effect of Low Temperature at Booting Stage on Photosynthetic System of Different Rice Materials in Cold Region

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