核桃晚霜危害特性及影响机制研究进展

doi:10.13304/j.nykjdb.2022.0952

摘要/Abstract

摘要：

早春晚霜冻害的发生给我国核桃生产带来严峻的考验，严重影响核桃生长发育、产量及收益。为明确我国核桃晚霜危害特性及影响机制，综述了核桃受晚霜危害程度的影响因素及其在响应低温胁迫时生理生化指标变化和分子机制。分析发现，关于生理变化相关的研究集中在细胞膜系统（相对电导率）、抗氧化系统（丙二醛、超氧化物歧化酶、过氧化物酶及过氧化氢酶）和渗透调节物质（可溶性糖、可溶性蛋白质及脯氨酸）3个方面，JrPAL、JrJAZ、JrGRAS、JrICE1、JrCBF/JrDREB1、JrCOR、JrWRKY、JrDHN、JrGST和JrPUB等基因的表达量受到低温胁迫的诱导。对未来的研究方向进行了展望，为进一步挖掘核桃应对晚霜危害的响应机制及采取有效的生产措施提供了科学依据。

关键词: 核桃, 晚霜危害, 低温胁迫, 影响因素, 机理

Abstract:

The occurrence of early spring and late frost damage brought severe challenges to the production of walnuts in China， seriously affecting their growth， development， yield and revenue. In order to clarify the harmful characteristics and influencing mechanisms of late frost on walnuts in China， the influencing factors of the degree of damage caused by late frost on walnuts， as well as the changes in physiological and biochemical indicators and molecular mechanisms in response to low temperature stress were reviewed. It was found that research on physiological changes focused on the three aspects including cell membrane system （relative conductivity）， antioxidant system （malondialdehyde， superoxide dismutase， peroxidase and catalase） and osmoregulation substances （soluble sugars， soluble protein and proline）， in addition， the expression levels of genes such as JrPAL， JrJAZ， JrGRAS， JrICE1， JrCBF/JrDREB1， JrCOR， JrWRKY， JrDHN， JrGST， and JrPUB were induced by low temperature stress. And prospects were put for future research directions， which provided scientific basis for further exploring the response mechanism of walnuts to late frost damage and adopting effective production measures.

Key words: walnut, late frost damage, low-temperature stress, influence factor, mechanism

中图分类号:

S664.1

张绥林, 李洋, 李琰, 张赟齐, 齐建勋, 侯智霞. 核桃晚霜危害特性及影响机制研究进展[J]. 中国农业科技导报, 2024, 26(4): 18-26.

Suilin ZHANG, Yang LI, Yan LI, Yunqi ZHANG, Jianxun QI, Zhixia HOU. Research Progress on Harmful Characteristics and Mechanism of Walnut Late Frost[J]. Journal of Agricultural Science and Technology, 2024, 26(4): 18-26.

图/表 2

图1 低温胁迫下核桃生理生化变化趋势注：↑表示物质含量上升。

Fig. 1 Physiological and biochemical trends of walnut under low temperature stressNote： ↑ indicates an increase in substance content.

图2 核桃响应低温胁迫的模式注：↑和↓分别表示基因表达量上调和下调。

Fig. 2 Pattern diagram of walnut response to low temperature stressNote： ↑ and ↓ indicate the upregulation and dounregulation of gene expression levels， respectively.

参考文献 42

1	郗荣庭. 果树栽培学总论[M]. 北京:中国农业出版社, 1997:303.
2	蒋瑶, 魏海林, 高昌虎, 等. 湖南低山丘陵区薄壳山核桃的开花物候期观测及品种配置[J]. 南京林业大学学报(自然科学版), 2021, 45(1): 53-62.
	JIANG Y, WEI H L, GAO C H, et al.. Observation on flowering phenology and variety combination of Carya illinoinensis in low mountains and hills of Hunan province [J]. J. Nanjing For. Univ. (Nat. Sci.), 2021, 45(1): 53-62.
3	王一峰, 陈耀年, 王瀚, 等. 我国核桃晚霜冻害研究进展[J]. 农业与技术, 2020, 40(11): 49-51.
4	杨亚利, 郑合清, 张淑敏, 等. 2018和2020年铜川核桃晚霜冻害分析[J]. 陕西气象, 2022(3): 64-68.
5	乐呼, 雷晓英, 袁晓霞, 等. 蓝田县核桃春季霜冻特征及风险区划[J]. 现代农业科技, 2022(5): 163-165, 169.
6	武静, 贺奇, 史敏华, 等. 山西省核桃晚霜危害调查及防御措施研究[J]. 山西林业科技, 2016, 45(3): 28-30.
	WU J, HE Q, SHI M H, et al.. Study on the damage and preventive measures of late frost to Juglans regia in Shanxi province [J]. Shanxi For. Sci. Technol., 2016, 45(3): 28-30.
7	樊江斌, 张金龙, 许新平, 等. 陕西2013年核桃晚霜冻害发生情况调查与分析[J]. 西北林学院学报, 2014, 29(5): 120-124.
	PAN J B, ZHANG J L, XU X P, et al.. An investigation on frozen damage of walnut trees in Shaanxi province [J]. J. Northwest For. Univ., 2014, 29(5): 120-124.
8	王荣梅, 张晓琴, 刘姣, 等. 新疆喀什地区近50 a来霜冻变化特征及其对农业的影响[J]. 干旱气象, 2013, 31(2): 309-312.
	WANG R M, ZHANG X Q, LIU J, et al.. Frost variation and its influence on agricultural in Kashi of Xinjiang in recent 50 years [J]. J. Arid Meteorol., 2013, 31(2): 309-312.
9	刘杜玲, 张博勇, 彭少兵, 等. 早实核桃物候期观察与避晚霜品种的筛选[J]. 北方园艺, 2011(24): 14-17.
	LIU D L, ZHANG B Y, PENG S B, et al.. Observation on phenological period and varieties choice in avoiding the late frost damage of early-fruiting walnut culivars [J]. Northern Hortic., 2011(24): 14-17.
10	武彦霞, 田鑫, 李夏媛, 等. 山西省核桃晚霜冻害及预防补救措施[J]. 果树资源学报, 2021, 2(1): 51-52.
	WU Y X, TIAN X, LI X Y, et al.. Late frost damage of walnut and preventive and remedial measures in Shanxi province [J]. J. Fruit Resour., 2021, 2(1): 51-52.
11	王一峰, 赵淑玲, 庞世伟, 等. 2018年成县核桃晚霜冻害调查与分析[J]. 陕西林业科技, 2018, 46(3): 32-35.
	WANG Y F, ZHAO S L, PANG S W, et al.. Investigation on freezing damage to walnut trees in Cheng county [J]. Shaanxi For. Sci. Technol., 2018, 46(3): 32-35.
12	YADAV S K. Cold stress tolerance mechanisms in plants. a review [J]. Agron. Sustain. Dev., 2010, 30(3): 515-527.
13	常月梅, 李永琛, 高俊仙. 核桃资源的抗晚霜性能评价[J]. 果树资源学报, 2022, 3(5): 15-19.
	CHANG Y M, LI Y C, GAO J X. Evaluation of late frost resistance of walnut genetic resources [J]. J. Fruit Resour., 2022, 3(5): 15-19.
14	冀中锐, 张树振, 史根生, 等. 核桃物候期观察与避晚霜品种的筛选[J]. 落叶果树, 2021, 53(1): 13-16.
	JI Z R, ZHANG S Z, SHI G S, et al.. Observation on the phenophase of walnut and varieties selection for late frost avoidance [J]. Deciduous Fruits, 2021, 53(1): 13-16.
15	王一峰, 赵淑玲, 王让军, 等. 甘肃成县核桃种质资源晚霜冻害调查[J]. 中国果树, 2020(1): 120-124.
16	仙米西努尔·克里木. 低温对新疆主要核桃品种枝条抗寒性生理指标的影响及抗冻措施[C]//中国环境科学学会2022年科学技术年会论文集(三).中国环境科学学会, 2022: 706-713.
17	张敏欢, 王建成, 杨红兰, 等. 新疆野核桃种质对低温胁迫的生理响应[J]. 应用生态学报, 2020, 31(8): 2558-2566.
	ZHANG M H, WANG J C, YANG H L, et al.. Physiological response of Xinjiang wild walnut germplasm to low temperature stress [J]. Chin. J. Appl. Ecol., 2020, 31(8): 2558-2566.
18	武彦霞, 史高川, 田鑫, 等. 5个核桃优系叶片抗寒性指标分析[J]. 果树资源学报, 2022, 3(6): 25-26, 37.
	WU Y X, SHI G C, TIAN X, et al.. Leaf cold resistance indexes analysis of five walnuts excellent individuals [J]. J. Fruit Resour., 2022, 3(6): 25-26, 37.
19	王一峰, 赵淑玲, 王瀚, 等. 不同核桃种质展叶期抗寒性的综合评价[J]. 经济林研究, 2019, 37(1): 50-60.
	WANG Y F, ZHAO S L, WANG H, et al.. Comprehensive evaluation on cold resistance of different walnut germplasms at leaf-expansion period [J]. Non-wood For. Res., 2019, 37(1): 50-60.
20	吴硕, 贾彦丽, 智福军. 低温胁迫下核桃枝条抗寒性综合评价[J]. 林业与生态科学, 2020, 35(3): 314-319.
	WU S, JIA Y L, ZHI F J. Comprehensive evaluation of cold resistance of walnut branches under low temperature stress [J]. For. Ecol. Sci., 2020, 35(3): 314-319.
21	任俊杰, 赵爽, 苏彦苹, 等. 春季低温胁迫对核桃抗氧化酶指标的影响[J]. 西北农林科技大学学报(自然科学版), 2016, 44(3): 75-81.
	REN J J, ZHAO S, SU Y P, et al.. Effects of low temperature stress in spring on antioxidant indexes of walnuts [J]. J. Northwest A&F Univ. (Nat. Sci.), 2016, 44(3): 75-81.
22	相昆, 徐颖, 王新亮, 等. 低温胁迫对核桃枝条活性氧代谢的影响[J]. 江西农业学报, 2014, 26(1): 35-37.
	XIANG K, XU Y, WANG X L, et al.. Effects of low temperature stress on active oxygen metabolism in walnut branches [J]. Acta Agric. Jiangxi, 2014, 26(1): 35-37.
23	田景花, 王红霞, 张志华, 等. 低温逆境对不同核桃品种抗氧化系统及超微结构的影响[J]. 应用生态学报, 2015, 26(5): 1320-1326.
	TIAN J H, WANG H X, ZHANG Z H, et al.. Effects of chilling stress on antioxidant system and ultrastructure of walnut cultivars [J]. Chin. J. Appl. Ecol., 2015, 26(5): 1320-1326.
24	韩立群, 马凯, 李亚利, 等. 低温胁迫下美国核桃和中国新疆核桃抗寒性综合评价[J]. 新疆农业科学, 2017, 54(10): 1804-1812.
	HAN L Q, MA K, LI Y L, et al.. Comprehensive evaluation of cold resistance of American walnut and China Xinjiang walnut under low temperature stress [J]. Xinjiang Agric. Sci., 2017, 54(10): 1804-1812.
25	刘荣, 刘清国, 范建新, 等. 果树抗寒性生理生化及分子机理研究进展[J]. 分子植物育种, 2017, 15(5): 2028-2034.
	LIU R, LIU Q G, FAN J X, et al.. Researchadvances in the cold-resistance physiology biochemical and molecular mechanism in fruit crops [J]. Mol. Plant Breed., 2017, 15(5): 2028-2034.
26	YANG G Y, ZHANG W H, LIU Z X, et al.. Both JrWRKY2 and JrWRKY7 of Juglans regia mediate responses to abiotic stresses and abscisic acid through formation of homodimers and interaction [J]. Plant Biol., 2017, 19(2): 268-278.
27	YANG G Y, ZHANG W H, SUN Y D, et al.. Two novel WRKY genes from Juglans regia, JrWRKY6 and JrWRKY53, are involved in abscisic acid-dependent stress responses [J]. Biol. Plantarum, 2017, 61(4): 611-621.
28	XU F, DENG G, CHENG S, et al.. Molecular cloning, characterization and expression of the phenylalanine ammonia-lyase gene from Juglans regia [J]. Molecules, 2012, 17(7): 7810-7823.
29	刘凯, 李津津, 王红霞, 等. 核桃JrJAZ基因家族鉴定与转录表达分析[J]. 河北农业大学学报, 2021, 44(1): 41-50.
	LIU K, LI J J, WANG H X, et al.. Identification and transcriptional expression analysis of walnut JrJAZ gene family [J]. J. Hebei Agric. Univ., 2021, 44(1): 41-50.
30	QUAN S W, NIU J X, ZHOU L, et al.. Genome-wide identification, classification, expression and duplication analysis of GRAS family genes in Juglans regia L. [J/OL]. Sci. Rep., 2019, 9(1): 11643[2022-10-16]. .
31	刘凯, 赵星哲, 王红霞, 等. 核桃JrGRAS基因家族鉴定与响应低温胁迫基因筛选[J]. 河北农业大学学报, 2021, 44(2): 27-40.
	LIU K, ZHAO X Z, WANG H X, et al.. Identification of walnut JrGRAS gene family and screening of genes in responce to low temperature stress [J]. J. Hebei Agric. Univ., 2021, 44(2): 27-40.
32	赵云. 喷施GA₃和ETH对核桃休眠芽萌发及抗寒力的影响[D]. 太原:山西农业大学, 2016.
	ZHAO Y. Effects of GA₃ and ETH on bud germinate and cold resistance for dormant buds of walnut [D]. Taiyuan:Shanxi Agricultural University, 2016.
33	TANG K, ZHAO L, REN Y Y, et al.. The transcription factor ICE1 functions in cold stress response by binding to the promoters of CBF and COR genes [J]. J. Integrat. Plant Biol., 2020, 62(3): 258-263.
34	ZUO Z F, KANG H G, PARK M Y, et al.. Overexpression of ICE1, a regulator of cold-induced transcriptome, confers cold tolerance to transgenic Zoysia japonica [J]. J. Plant Biol., 2019, 62(2): 137-146.
35	李津津, 刘凯, 王芳, 等. 核桃响应低温胁迫转录因子ICE1基因的克隆与分析[J]. 林业与生态科学, 2021, 36(3): 293-300.
	LI J J, LIU K, WANG F, et al.. Cloning and analysis of cold stress transcription factor ICE1 gene in Juglans regia [J]. For. Ecol. Sci., 2021, 36(3): 293-300.
36	徐丽, 陈新, 张力思, 等. 核桃JrCOR基因的克隆及表达分析[J]. 山东农业科学, 2018, 50(10): 13-18.
	XU L, CHEN X, ZHANG L S, et al.. Cloning and expression of JrCOR gene from walnut [J]. Shandong Agric. Sci., 2018, 50(10): 13-18.
37	RUBIO S, NORIEGA X, PEREZ F J. Abscisic acid (ABA) and low temperatures synergistically increase the expression of CBF/DREB1 transcription factors and cold-hardiness in grapevine dormant buds [J]. Ann. Bot., 2019, 123(4): 681-689.
38	徐丽, 陈新, 张力思, 等. 核桃JrCBF基因的克隆与表达和单核苷酸多态性分析[J]. 植物遗传资源学报, 2014, 15(2): 320-326.
	XU L, CHEN X, ZHANG L S, et al.. Molecular cloning, expression, and single nucleotide polymorphism analysis of transcription factor gene JrCBF in Juglans regia L. [J]. J. Plant Genet. Resour., 2014, 15(2): 320-326.
39	宋静武, 彭磊. 核桃CBF基因在低温胁迫中表达[J]. 湖北林业科技, 2019, 48(2): 7-13.
	SONG J W, PENG L. Investigating the expression of CBF gene in Juglans regia under cold stress [J]. Hubei For. Sci. Technol., 2019, 48(2): 7-13.
40	徐丽, 陈新, 魏海蓉, 等. 核桃Y₂SK₂型脱水素基因JrDHN的克隆、表达和单核苷酸多态性分析[J]. 园艺学报, 2014, 41(8): 1573-1582.
	XU L, CHEN X, WEI H R, et al.. Molecular cloning, expression, and single nucleotide polymorphism analysis of typical Y₂SK₂ dehydratin in Juglans [J]. Acta Hortic. Sin., 2014, 41(8): 1573-1582.
41	YANG G Y, XU Z G, PENG S B, et al.. In planta characterization of a tau class glutathione S-transferase gene from Juglans regia (JrGSTTau1) involved in chilling tolerance [J]. Plant Cell Rep., 2016, 35(3): 681-692.
42	姬新颖, 冯启科, 刘志月, 等. 核桃JrPUBs(U-box)基因家族鉴定与响应冷温胁迫基因筛选[J]. 河北农业大学学报, 2022, 45(6): 68-80.
	JI X Y, FENG Q K, LIU Z Y, et al.. Identification of walnut JrPUBs (U-box) gene family and screening of genes in responce to chilling stress [J]. J. Hebei Agric. Univ., 2022, 45(6): 68-80.

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