苹果抗寒性研究进展

doi:10.16420/j.issn.0513-353x.2025-0139

摘要/Abstract

摘要：

对近年来苹果抗寒性相关研究结果进行总结归纳，阐述了低温胁迫对苹果抗寒性的影响，比较了不同抗寒性的鉴定方法，列举了国内外苹果主要的抗寒品种和砧木，探究了苹果响应低温胁迫的生理及分子机制，最后指出了苹果抗寒性研究存在的问题及今后研究的主要方向，以期为进一步解析苹果抗寒机制及抗寒品种的选育提供参考。

关键词: 苹果, 抗寒, 机制, 进展

Abstract:

In this paper，the research results of apple cold resistance in recent years were summarized，the effects of low temperature stress on the cold resistance of apple were reviewed，the identification methods of different cold resistance were compared，the main cold-resistant apple varieties and rootstocks both domestically and internationally were listed，and the physiological and molecular mechanisms of apple in response to low temperature stress were explored. Finally，the problems existing in the study of apple cold resistance and the main direction of future research were pointed out. To provide support for further analysis of the cold resistance mechanism of apples and the breeding of cold resistance cultivars.

Key words: apple, cold resistance, mechanism, advance

徐功勋, 闫帅, 何美琦, 马怀宇, 吕德国, 秦嗣军, 程存刚, 赵德英. 苹果抗寒性研究进展[J]. 园艺学报, 2025, 52(8): 2021-2045.

XU Gongxun, YAN Shuai, HE Meiqi, MA Huaiyu, LÜ Deguo, QIN Sijun, CHENG Cungang, ZHAO Deying. Advances in the Research on Cold Resistance of Apple[J]. Acta Horticulturae Sinica, 2025, 52(8): 2021-2045.

图/表 5

参考文献 151

[1]	Abasi Kerimu, Meng Fanxue, Kudereti Rehanguli, Wang Jun, Li Huimin. 2023. Analysis of agro‐meteorological factors causing apple florescence freezing in zepu. Chinese Agricultural Science Bulletin, 39 (36):97-101. (in Chinese)
	克日木 · 阿巴司,孟凡雪,热汗古力 · 库得热提,汪俊,李慧民. 2023. 新疆泽普县苹果花期冻害农业气象因素分析. 中国农学通报, 39 (36):97-101.
[2]	An J P, Li R, Qu F J, You C X, Wang X F, Hao Y J. 2018. R2R3-MYB transcription factor MdMYB 23 is involved in the cold tolerance and proanthocyanidin accumulation in apple. The Plant Journal, 96 (3):562-577.
[3]	An J P, Liu Z Y, Zhang X W, Wang D R, Zeng F, You C X, Han Y. 2023. Brassinosteroid signaling regulator BIM1 integrates brassinolide and jasmonic acid signaling during cold tolerance in apple. Plant Physiology, 193 (2):1652-1674.
[4]	An J P, Wang X F, Zhang X W, You C X, Hao Y J. 2021. Apple B-box protein BBX37 regulates jasmonic acid mediated cold tolerance through the JAZ-BBX37-ICE1-CBF pathway and undergoes MIEL1-mediated ubiquitination and degradation. New Phytologist, 229 (5):2707-2729.
[5]	An J P, Xu R R, Liu X, Su L, Yang K, Wang X F, Wang G L, You C X. 2022. Abscisic acid insensitive 4 interacts with ICE1 and JAZ proteins to regulate ABA signaling-mediated cold tolerance in apple. Journal of Experimental Botany, 73 (3):980-997.
[6]	An J P, Yao J F, Wang X N, You C X, Wang X F, Hao Y J. 2017. MdHY 5 positively regulates cold tolerance via CBF-dependent and CBF-independent pathways in apple. Journal of Plant Physiology,218:275-281.
[7]	Bai Qinfeng, Huo Zhiguo, Wang Jinghong, Zhang Yong. 2019. Progress in research on meteorological disaster indicators of major fruit trees in China. Journal of Fruit Science, 36 (9):1229-1243. (in Chinese)
	柏秦凤, 霍治国, 王景红, 张勇. 2019. 中国主要果树气象灾害指标研究进展. 果树学报, 36 (9):1229-1243.
[8]	Bao Junhong, Bao Aomin, Yang Rong, Li Jinpu, Wang Baoxia, Zheng Dongsheng, He Yanhong. 2024. Comprehensive evaluation of cold resistance of apple Longfeng,Yueyan and their two hybrid strains in cold region. Journal of Fruit Science, 41 (2):241-251. (in Chinese)
	包俊宏, 包敖民, 杨荣, 李今普, 王宝侠, 郑东生, 何炎红. 2024. 寒地苹果龙丰、岳艳及其两个杂交优系抗寒性综合评价. 果树学报, 41 (2):241-251.
[9]	Beauvieux R, Wenden B, Dirlewanger E. 2018. Bud dormancy in perennial fruit tree species:a pivotal role for oxidative cues. Frontiers in Plant Science,9:657.
[10]	Bu Haidong, Gu Guangjun, Liu Chang, Hu Yinghui, Yang Yue, Yu Wenquan. 2023a. Breeding of a new cold-resistant apple cultivar ‘Longshuo’. Northern Horticulture,(18):157-160,2. (in Chinese)
	卜海东, 顾广军, 刘畅, 胡颖慧, 杨悦, 于文全. 2023a. 抗寒苹果新品种‘龙硕’的选育. 北方园艺,(18):157-160,2.
[11]	Bu Haidong, Gu Guangjun, Liu Chang, Hu Yinghui, Yang Yue, Cheng Xianmin, Yu Wenquan, Zhang Wujie, Gao Hongna, Lin Cunxue, Yang Xiaohua, Dong Hang. 2023b. A new apple cultivar‘Longyan’. Acta Horticulturae Sinica, 50 (S2):1-2. (in Chinese)
	卜海东, 顾广军, 刘畅, 胡颖慧, 杨悦, 程显敏, 于文全, 张武杰, 高洪娜, 林存学, 杨晓华, 董航. 2023b. 苹果新品种‘龙艳’. 园艺学报, 50 (S2):1-2.
[12]	Cai Y, Gao X, Mao J, Liu Y, Tong L, Chen X, Liu Y, Kou W, Chang C, Foster T, Yao J, Cornille A, Tahir M M, Liu Z, Yan Z, Lin S, Ma F, Ma J, Xing L, An N, Zuo X, Lv Y, Zhao Z, Li W, Li Q, Zhao C, Hu Y, Liu H, Wang C, Shi X, Ma D, Fei Z, Jiang Y, Zhang D. 2024. Genome sequencing of‘Fuji’apple clonal varieties reveals genetic mechanism of the spur-type morphology. Nature Communications, 15 (1):10082.
[122]	Wu Kun. 1992. New apple variety Longfeng. Modernizing Agriculture,(12):13. (in Chinese)
	吴昆. 1992. 苹果新品种龙丰. 现代化农业,(12):13.
[123]	Xie Y, Chen P, Yan Y, Bao C, Li X, Wang L, Shen X, Li H, Liu X, Niu C, Zhu C, Fang N, Shao Y, Zhao T, Yu J, Zhu J, Xu L, van Nocker S, Ma F, Guan Q. 2018. An atypical R2R3 MYB transcription factor increases cold hardiness by CBF-dependent and CBF-independent pathways in apple. New Phytologist, 218 (1):201-218.
[124]	Xu G, He M, Yan S, Lyu D, Cheng C, Zhao D, Qin S. 2025. Galactinol synthase gene 5(MdGolS5)enhances the cold resistance of apples by promoting raffinose family oligosaccharide accumulation. Plant Physiology and Biochemistry,220:109416.
[125]	Xu G, He M, Zhao D, Lyu D, Qin S. 2023a. Physiological and structural changes in apple tree branches of different varieties during dormancy. Horticulturae, 9 (8):947.
[126]	Xu G, Li L, Zhou J, He M, Lyu D, Zhao D, Qin S. 2023b. Integrated transcriptomics and metabolomics analyses reveal key genes and essential metabolic pathways for the acquisition of cold tolerance during dormancy in apple. Environmental and Experimental Botany,105413.
[127]	Xu G, Li L, Zhou J, Lyu D, Zhao D, Qin S. 2023c. Comparison of transcriptome and metabolome analysis revealed differences in cold resistant metabolic pathways in different apple cultivars under low temperature stress. Horticultural Plant Journal, 9 (2):183-198.
[13]	Chen D, Shao Q, Yin L, Younis A, Zheng B. 2019. Polyamine function in plants:metabolism,regulation on development,and roles in abiotic stress response. Frontiers in Plant Science,9:1945.
[14]	Chen X, Li S, Zhang D, Han M, Jin X, Zhao C, Wang S, Xing L, Ma J, Ji J, An N. 2019. Sequencing of a wild apple(Malus baccata) genome unravels the differences between cultivated and wild apple species regarding disease resistance and cold tolerance. Genes Genomes Genetics, 9 (7):2051-2060.
[15]	Chen Xuesen, Wang Nan, Peng Futian, Mao Zhiquan, Yin Chengmiao, Jiang Yuanmao, Ge Shunfeng, Hu Dagang, Li Yuanyuan, Du Yuanpeng, Yao Yuxin, Zhang Zongying. 2024. Advances in quality and maturity breeding of important deciduous fruit trees in China. Acta Horticulturae Sinica, 51 (1):8-26. (in Chinese)
	陈学森, 王楠, 彭福田, 毛志泉, 尹承苗, 姜远茂, 葛顺峰, 胡大刚, 李媛媛, 杜远鹏, 姚玉新, 张宗营. 2024. 中国重要落叶果树果实品质和熟期育种研究进展. 园艺学报, 51 (1):8-26.
[16]	Cheng Cungang, Cong Peihua, Man Shuduo, Wang Qiang, Kang Guodong. 2004. A new apple variety-Huajin. Acta Horticulturae Sinica,(6):829. (in Chinese)
	程存刚, 丛佩华, 满书铎, 王强, 康国栋. 2004. 苹果新品种——华金. 园艺学报,(6):829.
[17]	Cheng Cungang, Zhao Deying. 2019. Development orientation and trend of the apple industry under the new situation. China Fruits,(1):1-7. (in Chinese)
	程存刚, 赵德英. 2019. 新形势下我国苹果产业的发展定位与趋势. 中国果树,(1):1-7.
[18]	Cooke J E, Eriksson M E, Junttila O. 2012. The dynamic nature of bud dormancy in trees:environmental control and molecular mechanisms. Plant Cell Environment, 35 (10):1707-1728.
[19]	Daccord N, Celton J M, Linsmith G, Becker C, Choisne N, Schijlen E, van de Geest H, Bianco L, Micheletti D, Velasco R, Di Pierro E A, Gouzy J, Rees D J G, Guérif P, Muranty H, Durel C E, Laurens F, Lespinasse Y, Gaillard S, Aubourg S, Quesneville H, Weigel D, van de Weg E, Troggio M, Bucher E. 2017. High-quality de novo assembly of the apple genome and methylome dynamics of early fruit development. Nature Genetics, 49 (7):1099-1106.
[128]	Xu Gongxun, Zhou Jia, Lü Deguo, Qin Sijun. 2023c. Cold resistance evaluation of four apple varieties. Journal of Fruit Science, 40 (4):669-679. (in Chinese)
	徐功勋, 周佳, 吕德国, 秦嗣军. 2023. 4个苹果品种的抗寒性评价. 果树学报, 40 (4):669-679.
[129]	Xu H, Wang N, Wang Y, Jiang S, Fang H, Zhang J, Su M, Zuo W, Xu L, Zhang Z, Chen X. 2018. Overexpression of the transcription factor MdbHLH 33 increases cold tolerance of transgenic apple callus. Plant Cell,Tissue and Organ Culture,134:131-140.
[130]	Xu P Y, Xu L, Xu H F, He X W, He P, Chang Y S, Wang S, Zheng W Y, Wang C Z, Chen X, Li L G, Wang H B. 2023d. MdWRKY 40 is directly promotes anthocyanin accumulation and blocks MdMYB15L,the repressor of MdCBF2,which improves cold tolerance in apple. Journal of Integrative Agriculture, 22 (6):1704-1719.
[131]	Yan Zhongye, Yi Kai, Liu Zhi, Wang Dongmei, Lü Tianxing, Zhang Jing’e, Yang Feng. 2014. Breeding of new apple variety‘Wangxianghong’with high quality and late ripening. China Fruits,(3):4-6. (in Chinese)
	闫忠业, 伊凯, 刘志, 王冬梅, 吕天星, 张景娥, 杨锋. 2014. 苹果优质晚熟新品种‘望香红’的选育. 中国果树,(3):4-6.
[132]	Yang J, Guo X, Mei Q, Qiu L, Chen P, Li W, Mao K, Ma F. 2023. MdbHLH 4 negatively regulates apple cold tolerance by inhibiting MdCBF1/3 expression and promoting MdCAX3L-2 expression. Plant Physiology, 191 (1):789-806.
[133]	Yuan Jicun, Cheng Cungang, Zhao Deying, Liu Shangtao, Li Enmao. 2021. Effects of different interstocks on the growth,yield,and fruit quality of Hanfu apple. Chinese Journal of Applied Ecology, 32 (9):3145-3151. (in Chinese)
	袁继存, 程存刚, 赵德英, 刘尚涛, 厉恩茂. 2021. 不同中间砧木对寒富苹果生长、产量和果实品质的影响. 应用生态学报, 32 (9):3145-3151.
[134]	Yu Wenquan, Guo Jinhong, Gu Guangjun, Bu Haidong, Liu Chang, Feng Zhangli, Cheng Xianmin, Lin Mingji, Zhou Jinhua, Dong Xuemei, Xing Xing. 2017. A new apple cultivar‘Longshuai’. Acta Horticulturae Sinica, 44 (1):193-194. (in Chinese)
	于文全, 郭金洪, 顾广军, 卜海东, 刘畅, 冯章丽, 程显敏, 林明极, 周进华, 董雪梅, 邢星. 2017. 苹果新品种‘龙帅’. 园艺学报, 44 (1):193-194.
[135]	Zhang Bingbing, Li Yuebo, Song Hongwei, Zhao Chenhui. 2011. Breeding of a new cold-resistant dwarf rootstock GM310 for apple. China Fruits,(6):4-5,81. (in Chinese)
	张冰冰, 李粤渤, 宋洪伟, 赵晨辉. 2011. 苹果抗寒矮化砧木新品种GM310的选育. 中国果树,(6):4-5,81.
[136]	Zhang Bo, Liu Liqiang, Qin Wei, Wurenqimige. 2021. Study on physiological and biochemical mechanism of cold resistance of Malus sieversii. Non-wood Forest Research, 39 (4):60-68. (in Chinese)
	张博, 刘立强, 秦伟, 乌仁其米格. 2021. 新疆野苹果抗寒生理生化机制研究. 经济林研究, 39 (4):60-68.
[137]	Zhang Jing’e, Yi Kai, Liu Zhi, Wang Dongmei, Yang Feng, Yan Zhongye, Liu Yanjie. 2007. Qiyuexian,a hardy early season dual apple cultivar. Journal of Fruit Science,(6):865-866,728. (in Chinese)
	张景娥, 伊凯, 刘志, 王冬梅, 杨峰, 闫忠业, 刘延杰. 2007. 苹果抗寒早熟鲜食与加工兼用新品种——七月鲜的选育. 果树学报,(6):865-866,728.
[138]	Zhang L, Hu J, Han X, Li J, Gao Y, Richards C M, Zhang C, Tian Y, Liu G, Gul H, Wang D, Tian Y, Yang C, Meng M, Yuan G, Kang G, Wu Y, Wang K, Zhang H, Wang D, Cong P. 2019. A high-quality apple genome assembly reveals the association of a retrotransposon and red fruit colour. Nature Communications, 10 (1):1494.
[139]	Zhang Q, Yu J, Wang J, Hu D, Hao Y. 2017. Functional characterization of MdMYB 73 reveals its involvement in cold stress response in apple calli and Arabidopsis. Journal of Integrative Agriculture, 16 (10):2215-2221.
[20]	Devireddy A R, Zandalinas SI, Fichman Y, Mittler R. 2021. Integration of reactive oxygen species and hormone signaling during abiotic stress. The Plant Journal, 105 (2):459-476.
[21]	Ding Y, Shi Y, Yang S. 2019. Advances and challenges in uncovering cold tolerance regulatory mechanisms in plants. New Phytologist, 222 (4):1690-1704.
[22]	Ding Y, Shi Y, Yang S. 2020. Molecular regulation of plant responses to environmental temperatures. Molecular Plant, 13 (4):544-564.
[23]	Dong H, Wu C, Luo C, Wei M, Qu S, Wang S. 2020. Overexpression of MdCPK1a gene,a calcium dependent protein kinase in apple,increase tobacco cold tolerance via scavenging ROS accumulation. PLoS One, 15 (11):e0242139.
[24]	Duan N, Bai Y, Sun H, Wang N, Ma Y, Li M, Wang X, Jiao C, Legall N, Mao L, Wan S, Wang K, He T, Feng S, Zhang Z, Mao Z, Shen X, Chen X, Jiang Y, Wu S, Yin C, Ge S, Yang L, Jiang S, Xu H, Liu J, Wang D, Qu C, Wang Y, Zuo W, Xiang L, Liu C, Zhang D, Gao Y, Xu Y, Xu K, Chao T, Fazio G, Shu H, Zhong G Y, Cheng L, Fei Z, Chen X. 2017. Genome re-sequencing reveals the history of apple and supports a two-stage model for fruit enlargement. Nature Communications, 8 (1):249.
[25]	Fadón E, Fernandez E, Behn H, Luedeling E. 2020. A conceptual framework for winter dormancy in deciduous trees. Agronomy,10:241.
[26]	Fan Xiufang, Liu Xufeng, Zhang Jiantang, Shi Lianrang. 2005. The performance of SX126 apple dwarf rootstock in Shaanxi regional trial. Journal of Northwest A & F University(Natural Science Edition),(10):47-50. (in Chinese)
	樊秀芳, 刘旭峰, 张建堂, 史联让. 2005. SX126苹果矮化砧在陕西区试表现. 西北农林科技大学学报(自然科学版),(10):47-50.
[27]	Gao Dan. 2013. Breeding of a new cold-resistant apple cultivar‘Jin Ping’. China Fruits,(2):1-2,77. (in Chinese)
	高丹. 2013. 苹果抗寒新品种‘金苹’的选育. 中国果树,(2):1-2,77.
[28]	Gong Z, Xiong L, Shi H, Yang S, Herrera-Estrella L R, Xu G, Chao D Y, Li J, Wang P Y, Qin F, Li J, Ding Y, Shi Y, Wang Y, Yang Y, Guo Y, Zhu J K. 2020. Plant abiotic stress response and nutrient use efficiency. Science China. Life Sciences, 63 (5),635-674.
[29]	González N C, Delgado A, Else M, Hadley P. 2023. Apple(Malus × domestica Borkh.)dormancy-a review of regulatory mechanisms and agroclimatic requirements. Frontiers in Horticulture,2:1217689.
[30]	Guo M, Wang S, Liu H, Yao S, Yan J, Wang C, Miao B, Guo J, Ma F, Guan Q, Xu J. 2023. Histone deacetylase MdHDA 6 is an antagonist in regulation of transcription factor MdTCP15 to promote cold tolerance in apple. Plant Biotechnology Journal, 21 (11):2254-2272.
[31]	Han Zhenhai, Wang Yi, Zhang Xinzhong, Xu Xuefeng, Sun Yangwu, Shen Jun. 2013. Apple rootstock new variety Chistock #1. Journal of Agricultural Biotechnology, 21 (7):879-882. (in Chinese)
	韩振海, 王忆, 张新忠, 许雪峰, 孙扬吾, 沈隽. 2013. 苹果砧木新品种中砧1号. 农业生物技术学报, 21 (7):879-882.
[32]	Han Zhengwei, Guo Jing, Zhang Zhenfei, Pan Yue, Yang Xinrong. 2023. Breeding report of Xinjiang native apple Maping 1. Journal of Fruit Science, 40 (3):588-591. (in Chinese)
	韩政伟, 郭靖, 张振飞, 潘越, 杨新荣. 2023. 新疆乡土苹果新品种玛平1号的选育. 果树学报, 40 (3):588-591.
[33]	He M, Zhou J, Lyu D, Xu G, Qin S. 2024. Exogenous spermidine alleviated low-temperature damage by affecting polyamine metabolism and antioxidant levels in apples. Plants, 13 (8):1100.
[34]	Jia Y, Ding Y, Shi Y, Zhang X, Gong Z, Yang S. 2016. The cbfs triple mutants reveal the essential functions of CBFs in cold acclimation and allow the definition of CBF regulons in Arabidopsis. New Phytologist, 212 (2):345-353.
[35]	Jiang H, Zhou L J, Gao H N, Wang X F, Li Z W, Li Y Y. 2022. The transcription factor MdMYB 2 influences cold tolerance and anthocyanin accumulation by activating SUMO E3 ligase MdSIZ1 in apple. Plant Physiology, 189 (4):2044-2060.
[36]	Jiao Kuibao, Li Yuze, Song Penghui, Guo Baitao, Zhang Lei, Tang Ke, Wu Yuxi, Song Yanan. 2024. A new cold resistant small apple cultivar‘Zhenxing 1’. Acta Horticulturae Sinica, 51 (S2):25-26. (in Chinese)
	焦奎宝, 李雨泽, 宋鹏慧, 郭佰涛, 张磊, 唐克, 吴雨蹊, 宋亚楠. 2024. 抗寒小果型苹果新品种‘振兴1号’. 园艺学报, 51 (S2):25-26.
[37]	Jing Junli, Liu Mingxiao, Wei Xin, Xu Jizhong, Li Zhongyong, Zhang Xueying, Zhou Shasha. 2022. Evaluation of cold hardiness of several apple interstocks. Journal of Fruit Science, 39 (6):970-981. (in Chinese)
	井俊丽, 刘铭潇, 魏欣, 徐继忠, 李中勇, 张学英, 周莎莎. 2022. 几种苹果中间砧的抗寒性评价. 果树学报, 39 (6):970-981.
[38]	Jing Y, Pei T, Li C, Wang D, Wang Q, Chen Y, Li P, Liu C, Ma F. 2023a. Overexpression of the FERONIA receptor kinase MdMRLK2 enhances apple cold tolerance. The Plant Journal, 115 (1):236-252.
[39]	Jing J, Liu M, Yin B, Liang B, Li Z, Zhang X, Xu J, Zhou S. 2023b. Effects of 10 dwarfing interstocks on cold resistance of‘Tianhong 2’apple. Horticulturae, 9 (7):827.
[40]	Kan Zhiyong, Zhang Dehui, Li Zhongxing, Yu Sisi, Qian Qian, Fan Tianle, Li Xuewei, Ma Fengwang, Guan Qingmei. 2023. Evaluation of cold tolerance of 90 apple cultivars and genome wide association analysis. Acta Horticulturae Sinica, 50 (5):921-932. (in Chinese)
	坎智勇, 张德辉, 李中兴, 余思思, 钱谦, 樊天乐, 李雪薇, 马锋旺, 管清美. 2023. 90个苹果品种耐寒性评价和全基因组关联分析. 园艺学报, 50 (5):921-932.
[41]	Khan A, Carey SB, Serrano A, Zhang H, Hargarten H, Hale H, Harkess A, Honaas L. 2022. A phased,chromosome-scale genome of‘Honeycrisp’apple(Malus domestica). GigaByte,2022:gigabyte69.
[42]	Kidokoro S, Shinozaki K, Yamaguchi‐Shinozaki K. 2022. Transcriptional regulatory network of plant cold-stress responses. Trends in Plant Science, 27 (9):922-935.
[43]	Li B, Qu S, Kang J, Peng Y, Yang N, Ma B, Ruan Y L, Ma F, Li M, Zhu L. 2024. The MdCBF1/2-MdTST1/ 2 module regulates sugar accumulation in response to low temperature in apple. The Plant Journal, 118 (3):787-801.
[44]	Li Bowen, Jia Yanan, Zhou Qun, Wang Jiangbo. 2023. Evaluation of cold resistance of apple branches with different rootstock and spike combinations. Northern Horticulture,(19):36-43. (in Chinese)
	栗博文, 贾亚男, 周群, 王江波. 2023. 不同砧穗组合苹果枝条的抗寒性评价. 北方园艺,(19):36-43.
[45]	Li Ensheng, Rong Zhixiang, Li Xisen, Zhu Dan. 1990. Preliminary report on breeding of apple dwarfing rootstock excellent lines 77-33 and 77-34. Northern Fruits,( 3):24-25,27. (in Chinese)
	李恩生, 荣志祥, 李喜森, 朱丹. 1990. 苹果矮化砧木优系77-33、 77-34选育初报. 北方果树,(3):24-25,27.
[46]	Li Ge, Meng Xiaoqing, Cai Jing, Dong Tingting, Li Zongyun, Zhu Mingku. 2018. Advances in the function of reactive oxygen species in plant responses to abiotic stresses. Plant Physiology Journal, 54 (6):951-959. (in Chinese)
	李格, 孟小庆, 蔡敬, 董婷婷, 李宗芸, 朱明库. 2018. 活性氧在植物非生物胁迫响应中功能的研究进展. 植物生理学报, 54 (6):951-959.
[47]	Li Guoping, Liao Qingan, Ling Yizhang, Du Ping, Song Yuling. 2001. New apple cold-resistant variety-Xinping 4. China Fruits,(3):9-10. (in Chinese)
	李国萍, 廖庆安, 凌一章, 杜萍, 宋玉玲. 2001. 苹果抗寒新品种——新苹4号. 中国果树,(3):9-10.
[48]	Li Guoping, Ling Yizhang, Song Mei, Wang Shujuan, Song Yuling. 2004. Breeding of KM apple dwarfing rootstock. China Fruits,(4):5-7,2. (in Chinese)
	李国萍, 凌一章, 宋梅, 王淑娟, 宋玉玲. 2004. 苹果KM矮化砧的选育. 中国果树,(4):5-7,2.
[49]	Li Huaiyu, Liu Tao, Yu Zhihan. 2011. A new cold-resistant and high-quality apple cultivar‘Shenhong’. Northern Fruits,(1):56. (in Chinese)
	李怀玉, 刘涛, 于治汉. 2011. 抗寒优质苹果新品种‘沈红’. 北方果树,(1):56.
[50]	Li Huaiyu, Sun Hongxu, Qiao Fengqi, Cong Richun. 1995. A new cold-resistant and high-quality apple variety-Short Branch Hanfu. China Fruits,(1):1-2. (in Chinese)
	李怀玉, 孙红旭, 乔凤岐, 丛日春. 1995. 抗寒优质苹果新品种——短枝寒富. 中国果树,(1):1-2.
[51]	Li Shuxian, Teng Shuming, Guo Changcheng. 1987. A new cold-resistant apple variety-Longguan. China Fruits,(3):29-30. (in Chinese)
	李淑贤, 腾树明, 郭长城. 1987. 抗寒苹果新品种——龙冠. 中国果树,(3):29-30.
[52]	Li Tianzhong, Wang Baoan, Li Wei, Jiang Feng, Liu Zhi, Lü Deguo, Song Guozhu, Wang Shengyuan, Zhang Zhitao. 2021. SNP molecular markers for identifying cold resistance of Hanfu apple hybrids and their application. Beijing:CN202010841126.7. (in Chinese)
	李天忠, 王宝安, 李威, 姜峰, 刘志, 吕德国, 宋国柱, 王圣元, 张志涛. 2021. 鉴定寒富苹果杂种后代抗寒性的SNP分子标记及其应用. 北京:CN202010841126.7
[53]	Li W, Chu C, Li H, Zhang H, Sun H, Wang S, Wang Z, Li Y, Foster T M, López-Girona E, Yu J, Li Y, Ma Y, Zhang K, Han Y, Zhou B, Fan X, Xiong Y, Deng C H, Wang Y, Xu X, Han Z. 2024. Near-gapless and haplotype-resolved apple genomes provide insights into the genetic basis of rootstock-induced dwarfing. Nature Genetics, 56 (3):505-516.
[54]	Li X, Kui L, Zhang J, Xie Y, Wang L, Yan Y, Wang N, Xu J, Li C, Wang W, van Nocker S, Dong Y, Ma F, Guan Q. 2016. Improved hybrid de novo genome assembly of domesticated apple(Malus× domestica).. Gigascience)， 5 (1):35.
[55]	Li X, Ma Z, Song Y, Shen W, Yue Q, Khan A, Tahir M M, Wang X, Malnoy M, Ma F, Bus V, Zhou S, Guan Q. 2023. Insights into the molecular mechanisms underlying responses of apple trees to abiotic stresses. Horticulture Research, 10 (8):uhad144.
[56]	Li Xingchao, Li Juncai, Zhou Yanqi, Liu Ning, Yu Xianghe, Yin Kai, Liu Chengxian. 2001. Investigation on freezing injury of fruit trees in Liaoning Province. Northern Fruits,(5):21-25. (in Chinese)
	李兴超, 李俊才, 周晏起, 刘宁, 郁香荷, 伊凯, 刘成先. 2001. 辽宁省果树冻害调查. 北方果树,(5):21-25.
[57]	Li Yuebo, Zhang Bingbing, Song Hongwei, Liang Yinghai, Zhao Chenhui, Bao Zhenlong. 2016. Breeding of a new cold-resistant dwarf rootstock variety‘ZM-2000’for apple. China Fruits,(3):55-56. (in Chinese)
	李粤渤, 张冰冰, 宋宏伟, 梁英海, 赵晨辉, 包振龙. 2016. 苹果抗寒矮化砧木新品种‘ZM-2000’的选育. 中国果树,(3):55-56.
[58]	Li Z, Wang L, He J, Li X, Hou N, Guo J, Niu C, Li C, Liu S, Xu J, Xie Y, Zhang D, Shen X, Lu L, Geng D, Chen P, Jiang L, Wang L, Li H, Malnoy M, Deng C, Zou Y, Li C, Zhan X, Dong Y, Notaguchi M, Ma F, Xu Q, Guan Q. 2022. Chromosome-scale reference genome provides insights into the genetic origin and grafting-mediated stress tolerance of Malus prunifolia. Plant Biotechnol Journal, 20 (6):1015-1017.
[59]	Liang Y, Wang S, Zhao C, Ma X, Zhao Y, Shao J, Li Y, Li H, Song H, Ma H, Li H, Zhang B, Zhang L. 2020. Transcriptional regulation of bark freezing tolerance in apple(Malus domestica Borkh.). Horticulture Research, 7 (1):205.
[60]	Liao Qingan, Ling Yizhang, Li Guoping, Han Qiqing, Bai Wenju. 1995. A new cold-resistant apple variety-Xinshuai. China Fruits,(1):3-4. (in Chinese)
	廖庆安, 凌一章, 李国萍, 韩其庆, 白文菊. 1995. 苹果抗寒新品种——新帅. 中国果树,(1):3-4.
[61]	Liao Qingan, Ling Yizhang, Li Guoping, Han Qiqing, Bai Wenju. 1992. New cold-resistant apple variety-Xinguan. China Fruits,(2):17,6. (in Chinese)
	廖庆安, 凌一章, 李国萍, 韩其庆, 白文菊. 1992. 苹果抗寒新品种——新冠. 中国果树,(2):17,6.
[62]	Liu Guocheng, Ma Huaiyu, Lü Deguo, Qin Sijun, Du Guodong, Wang He. 2012. Research on shoot cold tolerance of‘Hanfu’apple. Northern Horticulture,(14):35-37. (in Chinese)
	刘国成, 马怀宇, 吕德国, 秦嗣军, 杜国栋, 王贺. 2012. ‘寒富’苹果枝条低温适应性研究. 北方园艺,(14):35-37.
[63]	Liu T, Li Y, Shi Y, Ma J, Peng Y, Tian X, Zhang N, Ma F, Li C. 2024a. γ-Aminobutyric acid mediated by MdCBF3-MdGAD 1 mitigates low temperature damage in apple. International Journal of Biological Macromolecules,279:135331.
[64]	Liu X, Wei J, Li S, Li J, Cao H, Huang D, Zhang D, Zhang Z, Gao T, Zhang Y, Ma F, Li C. 2024b. MdHY 5 positively regulates cold tolerance in apple by integrating the auxin and abscisic acid pathways. New Phytologist, 246 (5):2155-2173.
[65]	Liu Xiaofeng, Cong Peihua, Wang Qiang. 2023. Breeding of a new medium-maturing apple variety Zhongpingbeiyan. China Fruits,(5):110-111,141. (in Chinese)
	刘肖烽, 丛佩华, 王强. 2023. 苹果中熟新品种中苹贝艳的选育. 中国果树,(5):110-111,141.
[66]	Liu Xinglu, Wang Hongping, Sun Wentai, Dong Tie, Niu Junqiang, Ma Ming. 2021. Cold resistance evaluation of the shoots of 5 apple rootstocks. Journal of Fruit Science, 38 (8):1264-1274. (in Chinese)
	刘兴禄, 王红平, 孙文泰, 董铁, 牛军强, 马明. 2021. 5个砧木苹果枝条的抗寒性评价. 果树学报, 38 (8):1264-1274.
[67]	Liu Y, Gao X H, Tong L, Liu M Z, Zhou X K, Tahir M M, Xing L B, Ma J J, An N, Zhao C P, Yao J L, Zhang D. 2022. Multi-omics analyses reveal MdMYB 10 hypermethylation being responsible for a bud sport of apple fruit color. Horticulture Research,9:uhac179.
[68]	Liu Yanjie, Li Shuxian, Guo Changcheng, Zhao Yanhua, Cheng Xianmin. 2002. Breeding of a new cold-resistant apple variety‘Longhong’. China Fruits,(6):5-7,64. (in Chinese)
	刘延杰, 李淑贤, 郭长城, 赵艳华, 程显敏. 2002. 苹果抗寒新品种‘龙红’的选育. 中国果树,(6):5-7,64.
[69]	Liu Zhi, Li Xisen, Yi Kai, Rong Zhixiang, Yang Feng, Yang Wei. 2004. Breeding report of apple dwarf rootstock‘Liaozhen2’. Journal of Fruit Science,(5):501-502,505. (in Chinese)
	刘志, 李喜森, 伊凯, 荣志祥, 杨锋, 杨巍. 2004. 苹果矮化砧木‘辽砧2号’选育. 果树学报,(5):501-502,505.
[70]	Liu Zhi, Wang Dongmei, Zhang Jing’e, Yan Zhongye, Lü Tianxing, Yang Feng, Yi Kai. 2010. A new late ripening apple cultivar‘Yueping’. Acta Horticulturae Sinica, 37 (10):1695-1696. (in Chinese)
	刘志, 王冬梅, 张景娥, 闫忠业, 吕天星, 杨锋, 伊凯. 2010. 晚熟苹果新品种‘岳苹’. 园艺学报, 37 (10):1695-1696.
[71]	Liu H, Lu C, Wang S, Ren F, Wang H. 2021. Climate warming extends growing season but not reproductive phase of terrestrial plants. Global Ecology and Biogeography,30:950-960.
[72]	Lu Chao, Cai Huacheng, Zhang Guoqiang, Yang Tingzhen, Wang Jinzheng. 2018. Investigation and countermeasures of apple flowering freezing injury in Shanxi Province. Deciduous Fruits, 50 (3):15-17. (in Chinese)
	路超, 蔡华成, 张国强, 杨廷桢, 王金政. 2018. 山西省苹果花期冻害调查及灾后对策. 落叶果树, 50 (3):15-17.
[73]	Lü Deguo. 2017. Apple industry in China’s cold climate zone-Taking‘Hanfu’as an example. Northern Fruits,(1):41-42.
	吕德国. 2017. 中国冷凉气候区的苹果产业——以‘寒富’为例. 北方果树,(1):41-42.
[74]	Lü Tianxing, Wang Dongmei, Yan Zhongye, Jiang Xiaojun, Yi Kai, Liu Zhi. 2016. Breeding report of a new late ripening apple cultivar ‘Yueguan’. Journal of Fruit Science, 33 (10):1321-1323. (in Chinese)
	吕天星, 王冬梅, 闫忠业, 姜孝军, 伊凯, 刘志. 2016. 晚熟苹果新品种‘岳冠’的选育. 果树学报, 33 (10):1321-1323.
[75]	Ma Fengwang. 2023. Reflections on the development of China’s apple industry-current situation,problems and solutions. Deciduous Fruits, 55 (4):1-4. (in Chinese)
	马锋旺. 2023. 中国苹果产业发展的思考——现状、问题与出路. 落叶果树, 55 (4):1-4.
[76]	Ma Huaiyu, Xu Tianwei, Lü Deguo. 2018. Investigation on flower bud freezing injury of‘Hanfu’apple after continuous low temperature in central and northern Liaoning. Northern Fruits,(3):50-51. (in Chinese)
	马怀宇, 徐田伟, 吕德国. 2018. 辽宁中北部地区‘寒富’苹果连续低温后花芽冻害调查. 北方果树,(3):50-51.
[77]	Manasa S L, Panigrahy M, Panigrahi K C S, Rout G R. 2022. Overview of cold stress regulation in plants. The Botanical Review, 88 (3):359-387.
[78]	Man Shuduo, Niu Jianzhe, Cong Peihua, Qin Helan. 1999. Breeding of a new late-maturing apple cultivar‘Huahong’. China Fruits,(1):13-14. (in Chinese)
	满书铎, 牛健哲, 丛佩华, 秦贺兰. 1999. 苹果晚熟新品种‘华红’的选育. 中国果树,(1):13-14.
[79]	Meng Qingyan, Wang Xiaohong, Ta Na, Yuan Shuming, Gong Zhongxu, Zhou Ruihua. 1997. Apple dwarf cold-resistant germplasm-Zhaaishandingzi. China Fruits,(3):13-14. (in Chinese)
	孟庆炎, 王晓红, 塔娜, 袁淑明, 弓仲旭, 周瑞华. 1997. 苹果属矮生抗寒种质——扎矮山定子. 中国果树,(3):13-14.
[80]	Mei C, Yang J, Mei Q, Jia D, Yan P, Feng B, Mamat A, Gong X, Guan Q, Mao K, Wang J, Ma F. 2023. MdNAC 104 positively regulates apple cold tolerance via CBF-dependent and CBF-independent pathways. Plant Biotechnology Journal, 21 (10):2057-2073.
[81]	Mishra N, Jiang C, Chen L, Paul A, Chatterjee A, Shen G. 2023. Achieving abiotic stress tolerance in plants through antioxidative defense mechanisms. Frontiers in Plant Science,14:1110622.
[82]	Muthuramalingam P, Shin H, Adarshan S, Jeyasri R, Priya A, Chen JT, Ramesh M. 2022. Molecular insights into freezing stress in peach based on multi-omics and biotechnology:an overview. Plants, 11 (6):812.
[83]	Pagter M, Arora R. 2013. Winter survival and deacclimation of perennials under warmer climate:physiological perspectives. Physiologia Plantarum, 147 (1):75-87.
[84]	Qin S, Xu G, He J, Li L, Ma H, Lyu D. 2023. A chromosome-scale genome assembly of Malus domestica,a multi-stress resistant apple variety. Genomics, 115 (3):110627.
[85]	Qiu Ninggang, Zhang Tianfeng, Zhou Zhongwen, Zhang Hongfen, Chen Fan, Song Yanan. 2024. Analysis of impact of freezing damage from april 28th to 29th,2023 on flowering period of apples in Longdong. Chinese Agricultural Science Bulletin, 40 (20):109-114. (in Chinese)
	邱宁刚, 张天峰, 周忠文, 张洪芬, 陈帆, 宋亚男. 2024. 2023年4月28—29日冻害对陇东苹果花期的影响分析. 中国农学通报, 40 (20):109-114.
[86]	Qu Zhenjiang, Zhou Guangsheng, Wei Qinping. 2016. Meteorological disaster index and risk assessment of frost injury during apple florescence. Journal of Applied Meteorolgical Science, 27 (4):385-395. (in Chinese)
	屈振江, 周广胜, 魏钦平. 2016. 苹果花期冻害气象指标和风险评估. 应用气象学报, 27 (4):385-395.
[87]	Qu Zhenjiang, Zhou Guangsheng. 2017. The risk assessment of winter injury in main apple-producing regions of China. Journal of Natural Resources, 32 (5):829-840. (in Chinese)
	屈振江, 周广胜. 2017. 中国产区苹果越冬冻害的风险评估. 自然资源学报, 32 (5):829-840.
[88]	Raza A, Charagh S, Abbas S, Hassan M U, Saeed F, Haider S, Sharif R, Anand A, Corpas F J, Jin W, Varshney R K. 2023. Assessment of proline function in higher plants under extreme temperatures. Plant Biology, 25 (3):379-395.
[89]	Rodrigo J. 2000. Spring frosts in deciduous fruit trees-morphological damage and flower hardines. Scientia Horticulturae, 85 (3):155-173.
[90]	Saini A K, Jahed K R, Neres D F, Wright R C, Sherif S M. 2025. Investigating frost response,rootstock-dependent cold tolerance,and floral bud mortality in apple cultivars through transcriptomic insights. Plant Stress,16:100829.
[91]	Salmon Y, Lintunen A, Dayet A, Chan T, Dewar R, Vesala T, Hölttä T. 2020. Leaf carbon and water status control stomatal and nonstomatal limitations of photosynthesis in trees. New Phytologist, 226 (3):690-703.
[92]	Sapkota S, Liu J, Islam M T, Sherif S M. 2021. Changes in reactive oxygen species,antioxidants and carbohydrate metabolism in relation to dormancy transition and bud break in apple(Malus× domestica Borkh)cultivars. Antioxidants, 10 (10):1549.
[93]	Sengupta S, Mukherjee S, Basak P, Majumder A L. 2015. Significance of galactinol and raffinose family oligosaccharide synthesis in plants. Frontiers in Plant Science,6:656.
[94]	Shao Jing, Li Yuebo, Bao Zhenlong, Guo Wenrong, Lin Hongcheng, Zhang Bingbing. 2024. Breeding report of a new high-cold-resistance apple cultivar Binghong. Journal of Fruit Science, 41 (9):1903-1906. (in Chinese)
	邵静, 李粤渤, 包振龙, 郭文荣, 林泓成, 张冰冰. 2024. 抗寒小苹果新品种冰红的选育. 果树学报, 41 (9):1903-1906.
[95]	Shen X, Ping Y, Bao C, Liu C, Tahir M M, Li X, Song Y, Xu W, Ma F, Guan Q. 2023a. Mdm-miR160-MdARF17-MdWRKY 33 module mediates freezing tolerance in apple. The Plant Journal, 114 (2):262-278.
[96]	Shen X, Song Y, Ping Y, He J, Xie Y, Ma F, Li X, Guan Q. 2023b. The RNA-binding protein MdHYL 1 modulates cold tolerance and disease resistance in apple. Plant Physiology, 192 (3):2143-2160.
[97]	Shi Xianju. 1988. Cold-resistant early fruit high-yield apple-Jinhong. Shanxi Fruits,(3):42. (in Chinese)
	施献举. 1988. 抗寒早果丰产苹果——金红. 山西果树,(3):42.
[98]	Shi You, Chen Shuying, Diao Yongqiang, Wu Songmei. 2020. Evaluation on cold resistance in eight apple varieties introduced into Yili valley. Non-wood Forest Research, 38 (2):147-153. (in Chinese)
	石游, 陈淑英, 刁永强, 吴松梅. 2020. 引入伊犁河谷的8个苹果品种抗寒性评价. 经济林研究, 38 (2):147-153.
[99]	Su Y, Yang X, Wang Y, Li J, Long Q, Cao S, Wang X, Liu Z, Huang S, Chen Z, Peng Y, Zhang F, Xue H, Cao X, Zhang M, Yisilam G, Chu Z, Gao Y, Zhou Y, Liu Z, Xiao H, Tian X. 2024. Phased telomere-to-telomere reference genome and pangenome reveal an expansion of resistance genes during apple domestication. Plant Physiology, 195 (4):2799-2814.
[100]	Sun Fenglan, Zhang Lianxi, Lin Shufang. 1998. A new cold-resistant and early-maturing apple cultivar‘Xiangyahuang’. China Fruits,(4):19. (in Chinese)
	孙凤兰, 张连喜, 林淑芳. 1998. 抗寒早熟苹果新品种‘象牙黄’. 中国果树,(4):19.
[101]	Sun Limei, Qü Jingyan, Wang Yan, Li Xingchao. 2019. Review of occurrence and prevention of fruit freezing injury in Liaoning Province. Northern Fruits,(6):1-5,25. (in Chinese)
	孙丽梅, 曲静艳, 王岩, 李兴超. 2019. 辽宁省果树冻害发生和预防的回顾. 北方果树,(6):1-5,25.
[102]	Sun X, Jiao C, Schwaninger H, Chao C T, Ma Y, Duan N, Khan A, Ban S, Xu K, Cheng L, Zhong G Y, Fei Z. 2020. Phased diploid genome assemblies and pan-genomes provide insights into the genetic history of apple domestication. Nature Genetics, 52 (12):1423-1432.
[140]	Zhang Xinyu, Xu Xuemei, Sun Aijun. 1998. Introduction of cold-resistant dwarfing rootstock GM256. Northern Horticulture,(2):71. (in Chinese)
	张新宇, 徐学梅, 孙爱军. 1998. 抗寒矮化砧GM256引种. 北方园艺,(2):71.
[141]	Zhao D, Yuan J, Xu K, Cheng C, Li H. 2016. Selection of morphological,physiological and biochemical indices:evaluating dwarfing apple interstocks in cold climate zones. New Zealand Journal of Crop and Horticultural Science, 44 (4):291-311.
[142]	Zhao Deying. 2024a. Apple dwarf rootstocks——M and MM lines commonly used at home and abroad. Applied Technology and Information for Fruit Tree,(6):21-22. (in Chinese)
	赵德英. 2024a. 国内外常用苹果矮化砧木——M系和MM系. 果树实用技术与信息,(6):21-22.
[143]	Zhao Deying. 2024b. Domestic and foreign commonly used apple dwarf rootstock——Canada series. Applied Technology and Information for Fruit Tree,(11):19,25. (in Chinese)
	赵德英. 2024b. 国内外常用苹果矮化砧木——加拿大系列. 果树实用技术与信息,(11):19,25.
[144]	Zhao Deying. 2024c. Apple dwarf rootstock——Pilnitz series commonly used at home and abroad. Applied Technology and Information for Fruit Tree,(12):12. (in Chinese)
	赵德英. 2024c. 国内外常用苹果矮化砧木——Pillnitz系列. 果树实用技术与信息,(12):12.
[145]	Zhao Deying. 2024d. Apple dwarf rootstock——B series commonly used at home and abroad. Applied Technology and Information for Fruit Tree,(9):16. (in Chinese)
	赵德英. 2024d. 国内外常用苹果矮化砧木——B系. 果树实用技术与信息,(9):16.
[146]	Zhao Deying. 2024e. Apple dwarf rootstock-P series commonly used at home and abroad. Applied Technology and Information for Fruit Tree,(8):29. (in Chinese)
	赵德英. 2024e. 国内外常用苹果矮化砧木——P系. 果树实用技术与信息,(8):29.
[147]	Zhao Deying. 2024f. Apple dwarf rootstock——G line commonly used at home and abroad. Applied Technology and Information for Fruit Tree,(7):21-22. (in Chinese)
	赵德英. 2024f. 国内外常用苹果矮化砧木——G系. 果树实用技术与信息,(7):21-22.
[148]	Zhao Deying, Yan Shuai, Xu Kai, Yuan Jicun, Zhou Jiangtao, Cheng Cungang, Zhao Liangliang, Zhang Shaoyu, Hou Guixue. 2022. Effects of dwarfing interstocks on morphology and carbohydrates and nitrogen nutrition of Yanfu No. 3 apple nursery trees. Journal of Fruit Science, 39 (12):2289-2300. (in Chinese)
	赵德英, 闫帅, 徐锴, 袁继存, 周江涛, 程存刚, 赵亮亮, 张少瑜, 侯桂学. 2022. 矮化中间砧对烟富3号苹果苗木形态与碳氮养分的影响. 果树学报, 39 (12):2289-2300.
[103]	Tian Y, Thrimawithana A, Ding T, Guo J, Gleave A, Chagné D, Ampomah-Dwamena C, Ireland H S, Schaffer R J, Luo Z, Wang M, An X, Wang D, Gao Y, Wang K, Zhang H, Zhang R, Zhou Z, Yan Z, Zhang L, Zhang C, Cong P, Deng C H, Yao J L. 2022. Transposon insertions regulate genome-wide allele-specific expression and underpin flower colour variations in apple(Malus spp.). Plant Biotechnology Journal, 20 (7):1285-1297.
[104]	Turkyilmaz Unal B, Mentis O, Akyol E. 2015. Effects of exogenous salicylic acid on antioxidant activity and proline accumulation in apple(Malus domestica L.). Horticulture,Environment,and Biotechnology, 56 (5):606-611.
[105]	Velasco R, Zharkikh A, Affourtit J, Dhingra A, Cestaro A, Kalyanaraman A, Fontana P, Bhatnagar S K, Troggio M, Pruss D, Salvi S, Pindo M, Baldi P, Castelletti S, Cavaiuolo M, Coppola G, Costa F, Cova V, Dal Ri A, Goremykin V, Komjanc M, Longhi S, Magnago P, Malacarne G, Malnoy M, Micheletti D, Moretto M, Perazzolli M, Si-Ammour A, Vezzulli S, Zini E, Eldredge G, Fitzgerald L M, Gutin N, Lanchbury J, Macalma T, Mitchell J T, Reid J, Wardell B, Kodira C, Chen Z, Desany B, Niazi F, Palmer M, Koepke T, Jiwan D, Schaeffer S, Krishnan V, Wu C, Chu V T, King S T, Vick J, Tao Q, Mraz A, Stormo A, Stormo K, Bogden R, Ederle D, Stella A, Vecchietti A, Kater M M, Masiero S, Lasserre P, Lespinasse Y, Allan A C, Bus V, Chagné D, Crowhurst R N, Gleave A P, Lavezzo E, Fawcett J A, Proost S, Rouzé P, Sterck L, Toppo S, Lazzari B, Hellens R P, Durel C E, Gutin A, Bumgarner R E, Gardiner S E, Skolnick M, Egholm M, Van de Peer Y, Salamini F, Viola R. 2010. The genome of the domesticated apple(Malus × domestica Borkh.). Nature Genetics, 42 (10):833-839.
[106]	Wang Baoxia, Dong Zhiyuan, Zhang Xiujuan, Bao Mulin, Fu Jianping. 2014. High quality crabapple——‘Saiwaihong’selective breeding. Journal of Fruit Science, 31 (4):739-741,524. (in Chinese)
	王宝侠, 董志源, 张秀君, 包木林, 付建平. 2014. 优质小苹果——‘塞外红’的选育. 果树学报, 31 (4):739-741,524.
[107]	Wang D R, Zhang X W, Xu R R, Wang G L, You C X, An J P. 2022. Apple U-box-type E 3 ubiquitin ligase MdPUB23 reduces cold-stress tolerance by degrading the cold-stress regulatory protein MdICE1. Horticulture Research,9:uhac171.
[108]	Wang Dongmei, Liu Zhi, Yi Kai, Zhang Jing’e, Yan Zhongye, Lü Tianxing. 2015. Breeding of a new late-maturing apple cultivar‘Yuehua’. China Fruits,(1):1-3,85. (in Chinese)
	王冬梅, 刘志, 伊凯, 张景娥, 闫忠业, 吕天星. 2015. 苹果晚熟新品种‘岳华’的选育. 中国果树,(1):1-3,85.
[109]	Wang Dongmei, Yi Kai, Liu Zhi, Lü Tianxing, Zhang Jing’e, Yan Zhongye. 2017. Breeding report of a new middle ripening apple cultivar‘Yueyan’. Journal of Fruit Science, 34 (4):515-518. (in Chinese)
	王冬梅, 伊凯, 刘志, 吕天星, 张景娥, 闫忠业. 2017. 中熟苹果新品种‘岳艳’的选育. 果树学报, 34 (4):515-518.
[110]	Wang Dongmei, Liu Zhi, Zhang Jing’e, Yi Kai, Yan Zhongye, Lü Tianxing. 2013. Breeding of a new mid-late maturing apple cultivar ‘Yueyanghong’. China Fruits,(3):1-3,85. (in Chinese)
	王冬梅, 刘志, 张景娥, 伊凯, 闫忠业, 吕天星. 2013. 苹果中晚熟新品种‘岳阳红’的选育. 中国果树,(3):1-3,85.
[111]	Wang Hongping, Dong Tie, Liu Xinglu, Yin Xiaoning, Sun Wentai, Niu Junqiang, Ma Ming. 2020. A study on the cold resistance and the semi-lethal temperatures for branches of five apple rootstock cultivars. Journal of Fruit Science, 37 (4):495-501. (in Chinese)
	王红平, 董铁, 刘兴禄, 尹晓宁, 孙文泰, 牛军强, 马明. 2020. 5个苹果砧木品种枝条的低温半致死温度及耐寒性评价. 果树学报, 37 (4):495-501.
[112]	Wang Jinghong, Bai Qinfeng, Liang Yi, Qu Zhenjiang, Zhang Tao, Zhang Yong. 2015. Study on freezing index and weather conditions causing Shaanxi apple florescence freezing injury in 2013. Journal of Fruit Science, 32 (1):100-107,174. (in Chinese)
	王景红, 柏秦凤, 梁轶, 屈振江, 张焘, 张勇. 2015. 2013年陕西苹果花期冻害气象条件分析及受冻指标研究. 果树学报, 32 (1):100-107,174.
[113]	Wang P, Liu W C, Han C, Wang S, Bai M Y, Song C P. 2024. Reactive oxygen species:Multidimensional regulators of plant adaptation to abiotic stress and development. Journal of Integrative Plant Biology, 66 (3):330-367.
[114]	Wang Qiang, Cong Peihua. 2018. Breeding of a new mid-late maturing apple cultivar‘Huaqing’. China Fruits,(5):73-74. (in Chinese)
	王强, 丛佩华. 2018. 苹果中晚熟新品种‘华庆’的选育. 中国果树,(5):73-74.
[115]	Wang Rongying, Li Chunqiang, Wu Yan, Li Xin. 2012. Impact of climate changes on freezing injury to fruit trees and defensive measures. Journal of Anhui Agricultural Sciences, 40 (9):5310-5312,5316. (in Chinese)
	王荣英, 李春强, 吴雁, 李新. 2012. 气候变化对果树冻害的影响及其防御对策. 安徽农业科学, 40 (9):5310-5312,5316.
[116]	Wang Y, Jiang H, Mao Z, Liu W, Jiang S, Xu H, Su M, Zhang J, Wang N, Zhang Z, Chen X. 2021. Ethylene increases the cold tolerance of apple via the MdERF1B-MdCIbHLH 1 regulatory module. The Plant Journal, 106 (2):379-393.
[117]	Wang Y, Wang J, Sarwar R, Zhang W, Geng R, Zhu K M, Tan X L. 2024. Research progress on the physiological response and molecular mechanism of cold response in plants. Frontiers in Plant Science,15:1334913.
[118]	Wang Yanxiu, Shi Xiaoyun, Cheng Li. 2018. Overwintering performance of 7 apple dwarfing interstocks in central Gansu irrigation area. Gansu Agricultural Science and Technology,(7):3-6. (in Chinese)
	王延秀, 石晓昀, 程丽. 2018. 7个苹果矮化中间砧在甘肃中部灌区的越冬表现. 甘肃农业科技,(7):3-6.
[119]	Wang Y X, Hu Y, Chen B H, Zhu Y F, Dawuda M M, Svetl S. 2018. Physiological mechanisms of resistance to cold stress associated with 10 elite apple rootstocks. Journal of Integrative Agriculture, 17 (4):857-866.
[120]	Wei Lu, Niu Zimian, Zhang Dandan, He Yongbo, Xie Peng, Liu Yang. 2024. A new cold resistant apple dwarf rootstock cultivar‘SC6’. Acta Horticulturae Sinica, 51 (S2):27-28. (in Chinese)
	蔚露, 牛自勉, 张丹丹, 何永波, 谢鹏, 刘阳. 2024. 抗寒苹果矮化砧木新品种‘SC6’. 园艺学报, 51 (S2):27-28.
[121]	Wu Liren, Liu Wendong, Liu Jinjiang, Liu Yanjie, Cheng Xianmin, Wang Lidong. 2013. Breeding and main traits performance of small apple‘Zixiang’. Northern Horticulture,(7):176-178. (in Chinese)
	吴立仁, 刘文东, 刘金江, 刘延杰, 程显敏, 王丽冬. 2013. 小苹果‘紫香’的选育及主要性状表现. 北方园艺,(7):176-178.
[149]	Zhou L J, Li Y Y, Zhang R F, Zhang C L, Xie X B, Zhao C, Hao Y J. 2017. The small ubiquitin-like modifier E 3 ligase MdSIZ1 promotes anthocyanin accumulation by sumoylating MdMYB1 under low-temperature conditions in apple. Plant Cell and Environment, 40 (10):2068-2080.
[150]	Zhou Jiaxu, Yu Nianwen, Li Chenghui, Li Hongjian, Zhang Xiumei, Liu Ying. 2024. Research progress on characteristics and utilization of apple rootstock resources in China. Northern Horticulture,(12):126-133. (in Chinese)
	周家旭, 于年文, 里程辉, 李宏建, 张秀美, 刘影. 2024. 我国苹果砧木资源特性及利用研究进展. 北方园艺,(12):126-133.
[151]	Zhuang Xingyue, Li Wensheng, Zhang Yinghui, Zeng Yuzhi, Zhang Zhenjun. 2024. Evaluation of cold resistance of 9 kinds of apple shoots combinedwith scion-rootstock in Aksu area. Non-wood Forest Research, 42 (1):274-283,303. (in Chinese)
	庄星月, 李文胜, 张颖慧, 曾玉智, 张振军. 2024. 阿克苏地区9种苹果砧穗组合枝条的抗寒性评价. 经济林研究, 42 (1):274-283,303.

类型 Type	品种 Cultivar	亲本（父 × 母） Parent（male × female）	选育单位 Breeding organization	平均单果质量/g Average fruit weight	可溶性固形物/% Total soluble solid	总酸含量/% Total acid content	成熟期 Mature period	参考文献 Reference
苹果 Apple	寒富 Hanfu	富士 × 东光 Fuji × Dongguang	沈阳农业大学 Shenyang Agricultural University	250.00	15.20	0.34	9月下旬（辽宁沈阳）Late September（Shenyang，Liaoning）	李怀玉等,1995
	华红 Huahong	惠 × 金冠 Megumi × Golden Delicious	中国农业科学院果树研究所 Research Institute of Pomology，Chinese Academy of Agricultural Sciences	185.00	15.10	0.48	10月中上旬（辽宁兴城）Early and middle October（Xingcheng，Liaoning）	满书铎等,1999
	华庆 Huaqing	华富 × 金冠 Huafu × Golden Delicious		223.00	15.50	0.50	9月下旬（辽宁兴城）Late September（Xingcheng，Liaoning）	王强和丛佩华,2018
	华金 Huajin	好矮生 × 金矮生 Well spur × Gold spur		250.00	14.40	0.35	9月下旬（辽宁兴城）Late September（Xingcheng，Liaoning）	程存刚等,2004
	岳冠 Yueguan	岳帅 × 寒富 Yueshuai × Hanfu	辽宁省果树科学研究所 Liaoning Institute of Pomology	225.00	15.40	0.39	9月下旬（辽宁熊岳）Late September（Xiongyue，Liaoning）	吕天星等,2016
	岳艳 Yueyan	珊夏 × 寒富 Sansa × Hanfu		240.00	13.40	0.42	9月上旬（辽宁熊岳）Early September（Xiongyue，Liaoning）	王冬梅等,2017
	岳苹 Yueping	岳帅 × 寒富 Yueshuai × Hanfu		295.00	15.30	0.22	10月中旬（辽宁熊岳）Mid-October （Xiongyue，Liaoning）	刘志等,2010
	岳华 Yuehua	岳帅 × 寒富 Yueshuai × Hanfu		215.00	15.50	0.37	10月中旬（辽宁熊岳）Mid-October （Xiongyue，Liaoning）	王冬梅等,2015
	望香红 Wangxianghong	富士与红星混栽苹果园，亲本不详 Apple orchard with mixed cultivation of Fuji and Red Star cultivars，parental origin unspecified		240.00	14.10	0.32	10月中旬（辽宁瓦房店）Mid-October（Wafangdian，Liaoning）	闫忠业等,2014
	岳阳红 Yueyang hong	东光 × 富士 Dongguang × Fuji		205.00	15.22	0.49	9月下旬（辽宁营口）Late September（Yingkou，Liaoning）	王冬梅等,2013
	新帅 Xinshuai	新冬 × 金冠 Xindong × Golden Delicious	新疆生产建设兵团第七师农业科学研究所Xinjiang Production and Construction Corps 7th Division Institute of Agricultural Sciences	216.00	13.30	0.35	9月下旬至10月中旬（新疆奎屯） Late September to mid-October （Kuitun，Xinjiang）	廖庆安等,1995
	新冠 Xinguan	新冬 × 金冠 Xindong × Golden Delicious		143.00	14.70		9月中旬（新疆奎屯） Mid-September （Kuitun，Xinjiang）	廖庆安等,1992
	新苹4号 Xinping 4	新光 × 金冠 Xinguang × Golden Delicious		210.00	14.70	0.36	9月上中旬（新疆奎屯） Early and middle September （Kuitun，Xinjiang）	李国萍等,2001
	玛平1号 Maping 1	不详 Parental origin unspecified	新疆林科院园林绿化研究所Research Institute of Landscaping，Xinjiang Academy of Forestry Sciences	200.00	14.50		9月下旬（新疆北疆地区） Late September（Northern Xinjiang）	韩政伟等,2023
类型 Type	品种 Cultivar	亲本（父 × 母） Parent（male × female）	选育单位 Breeding organization	平均单果质量/g Average fruit weight	可溶性固形物/% Total soluble solid	总酸含量/% Total acid content	成熟期 Mature period	参考文献 Reference
小苹果 Crab apple	沈红 Shenhong	富士 × 金红 Fuji × Jinhong	沈阳农业大学 Shenyang Agricultural University	155.00	16.80		9月上旬（辽宁沈阳） Early September（Shenyang，Liaoning）	李怀玉等,2011
	中苹贝艳 Zongping beiyan	金冠 × 龙丰 Golden Delicious × Longfeng	中国农业科学院果树研究所 Research Institute of Pomology，Chinese Academy of Agricultural Sciences	68.00	18.00	0.27	10月上旬（辽宁兴城） Early October（Xingcheng，Liaoning）	刘肖烽等,2023
	七月鲜 Qiyuexian	铃铛 × 佚名大苹果 Lingdang × Anonymous apple	辽宁省果树科学研究所Liaoning Institute of Pomology	50.70	13.94	1.22	7月下旬至8月中旬（辽宁熊岳） Late July to mid-August （Xiongyue，Liaoning）	张景娥等,2007
	金红 Jinhong	红太平 × 金冠 Hongtaiping × Golden Delicious	吉林省农业科学院果树研究所 Pomology Research Institute，Jilin Academy of Agricultural Sciences	70.00	12.00	0.70	9月上中旬（吉林公主岭） Early to mid-September （Gongzhuling，Jilin）	施献举,1988
	象牙黄 Xiangya- huang	蛟河1号 × 金红 Jiaohe No.1 × Jinhong		120.00	13.00	0.94	7月下旬至8月上旬（吉林公主岭） From late July to early August （Gongzhuling，Jilin）	孙凤兰等,1998
	冰红 Binghong	金冠 × 红海棠 Golden Delicious × Red crabapple		64.00	12.40	0.49	9月上中旬（吉林公主岭） Early to mid-September （Gongzhuling，Jilin）	邵静等,2024
	塞外红 Saiwai- hong	不详 Parental origin unspecified	通辽市林业和草原研究所Tongliao Forestry and Grass land Science Research Institute	58.30	17.31	0.87	9月中下旬（内蒙古通辽） Mid-late September（Tongliao，Inner Mongolia）	王宝侠等,2014
	金萍 Jinping	宁酥 × 金红 Ningsu × Jinhong	辽宁农业职业技术学院 Liaoning Agricultural Vocational and Technical College	110.00	18.25	0.28	9月中旬（辽宁铁岭） Mid-September（Tieling，Liaoning）	高丹,2013
	紫香 Zixiang	赤阳 × 红海棠 Chiyang × Red crabapple	黑龙江省农业科学院浆果研究所 Berry Research Institute，Heilongjiang Academy of Agricultural Sciences	45.00	12.70	0.46	8月下旬（黑龙江宝清） Late August（Baoqing，Heilongjiang）	吴立仁等,2013
	龙丰 Longfeng	大秋 × 金红 Daqiu × Jinhong	黑龙江省农业科学院牡丹江分院 Mudanjiang Branch，Heilongjiang Academy of Agricultural Sciences	55.10	14.70	3.88	9月中旬（黑龙江牡丹江）Mid-September（Mudanjiang，Heilongjiang）	吴昆,1992
	龙红 Longhong	富士 × 铃铛 Fuji × Lingdang		75.00	16.10	0.37	8月下旬（黑龙江牡丹江） Late August （Mudanjiang，Heilongjiang）	刘延杰等,2002
	龙帅 Longshuai	金红 × 寒富 Jinhong × Hanfu		119.00	13.90	0.56	9月中旬（黑龙江牡丹江）Mid-September（Mudanjiang，Heilongjiang）	于文全等,2017
	龙冠 Longguan	七月鲜 × 金冠 Qiyuexian × Golden Delicious		94.50	13.00	0.17	9月上旬（黑龙江牡丹江） Early September（Mudanjiang，Heilongjiang）	李淑贤等,1987
	龙硕 Longshuo	龙丰自然芽变 Longfeng natural bud mutation		110.60	14.00	1.20	9月中下旬（黑龙江牡丹江）Mid-late September（Mudanjiang，Heilongjiang）	卜海东等,2023a
	龙艳 Longyan	金红自然芽变 Jinhong natural bud mutation		91.00	15.10	0.81	9月上中旬（黑龙江牡丹江）Early and middle September（Mudanjiang，Heilongjiang）	卜海东等,2023b
类型 Type	品种 Cultivar	亲本（父 × 母） Parent（male × female）	选育单位 Breeding organization	平均单果质量/g Average fruit weight	可溶性固形物/% Total soluble solid	总酸含量/% Total acid content	成熟期 Mature period	参考文献 Reference
小苹果 Crab apple	振兴1号 Zhenxing 1	龙冠 × 冻果 Longguan × Dongguo	黑龙江省农业科学院乡村振兴科技研究所 Institute of Rural Revitalization Technology，Heilongjiang Academy of Agricultural Sciences	43.00	16.50	0.48	9月下旬（黑龙江哈尔滨）Late September（Harbin，Heilongjiang）	焦奎宝等,2024

类型 Type	品种 Cultivar	亲本（父 × 母） Parent（male × female）	选育单位 Breeding organization	平均单果质量/g Average fruit weight	可溶性固形物/% Total soluble solid	总酸含量/% Total acid content	成熟期 Mature period	参考文献 Reference
苹果 Apple	寒富 Hanfu	富士 × 东光 Fuji × Dongguang	沈阳农业大学 Shenyang Agricultural University	250.00	15.20	0.34	9月下旬（辽宁沈阳）Late September（Shenyang，Liaoning）	李怀玉等,1995
	华红 Huahong	惠 × 金冠 Megumi × Golden Delicious	中国农业科学院果树研究所 Research Institute of Pomology，Chinese Academy of Agricultural Sciences	185.00	15.10	0.48	10月中上旬（辽宁兴城）Early and middle October（Xingcheng，Liaoning）	满书铎等,1999
	华庆 Huaqing	华富 × 金冠 Huafu × Golden Delicious		223.00	15.50	0.50	9月下旬（辽宁兴城）Late September（Xingcheng，Liaoning）	王强和丛佩华,2018
	华金 Huajin	好矮生 × 金矮生 Well spur × Gold spur		250.00	14.40	0.35	9月下旬（辽宁兴城）Late September（Xingcheng，Liaoning）	程存刚等,2004
	岳冠 Yueguan	岳帅 × 寒富 Yueshuai × Hanfu	辽宁省果树科学研究所 Liaoning Institute of Pomology	225.00	15.40	0.39	9月下旬（辽宁熊岳）Late September（Xiongyue，Liaoning）	吕天星等,2016
	岳艳 Yueyan	珊夏 × 寒富 Sansa × Hanfu		240.00	13.40	0.42	9月上旬（辽宁熊岳）Early September（Xiongyue，Liaoning）	王冬梅等,2017
	岳苹 Yueping	岳帅 × 寒富 Yueshuai × Hanfu		295.00	15.30	0.22	10月中旬（辽宁熊岳）Mid-October （Xiongyue，Liaoning）	刘志等,2010
	岳华 Yuehua	岳帅 × 寒富 Yueshuai × Hanfu		215.00	15.50	0.37	10月中旬（辽宁熊岳）Mid-October （Xiongyue，Liaoning）	王冬梅等,2015
	望香红 Wangxianghong	富士与红星混栽苹果园，亲本不详 Apple orchard with mixed cultivation of Fuji and Red Star cultivars，parental origin unspecified		240.00	14.10	0.32	10月中旬（辽宁瓦房店）Mid-October（Wafangdian，Liaoning）	闫忠业等,2014
	岳阳红 Yueyang hong	东光 × 富士 Dongguang × Fuji		205.00	15.22	0.49	9月下旬（辽宁营口）Late September（Yingkou，Liaoning）	王冬梅等,2013
	新帅 Xinshuai	新冬 × 金冠 Xindong × Golden Delicious	新疆生产建设兵团第七师农业科学研究所Xinjiang Production and Construction Corps 7th Division Institute of Agricultural Sciences	216.00	13.30	0.35	9月下旬至10月中旬（新疆奎屯） Late September to mid-October （Kuitun，Xinjiang）	廖庆安等,1995
	新冠 Xinguan	新冬 × 金冠 Xindong × Golden Delicious		143.00	14.70		9月中旬（新疆奎屯） Mid-September （Kuitun，Xinjiang）	廖庆安等,1992
	新苹4号 Xinping 4	新光 × 金冠 Xinguang × Golden Delicious		210.00	14.70	0.36	9月上中旬（新疆奎屯） Early and middle September （Kuitun，Xinjiang）	李国萍等,2001
	玛平1号 Maping 1	不详 Parental origin unspecified	新疆林科院园林绿化研究所Research Institute of Landscaping，Xinjiang Academy of Forestry Sciences	200.00	14.50		9月下旬（新疆北疆地区） Late September（Northern Xinjiang）	韩政伟等,2023
类型 Type	品种 Cultivar	亲本（父 × 母） Parent（male × female）	选育单位 Breeding organization	平均单果质量/g Average fruit weight	可溶性固形物/% Total soluble solid	总酸含量/% Total acid content	成熟期 Mature period	参考文献 Reference
小苹果 Crab apple	沈红 Shenhong	富士 × 金红 Fuji × Jinhong	沈阳农业大学 Shenyang Agricultural University	155.00	16.80		9月上旬（辽宁沈阳） Early September（Shenyang，Liaoning）	李怀玉等,2011
	中苹贝艳 Zongping beiyan	金冠 × 龙丰 Golden Delicious × Longfeng	中国农业科学院果树研究所 Research Institute of Pomology，Chinese Academy of Agricultural Sciences	68.00	18.00	0.27	10月上旬（辽宁兴城） Early October（Xingcheng，Liaoning）	刘肖烽等,2023
	七月鲜 Qiyuexian	铃铛 × 佚名大苹果 Lingdang × Anonymous apple	辽宁省果树科学研究所Liaoning Institute of Pomology	50.70	13.94	1.22	7月下旬至8月中旬（辽宁熊岳） Late July to mid-August （Xiongyue，Liaoning）	张景娥等,2007
	金红 Jinhong	红太平 × 金冠 Hongtaiping × Golden Delicious	吉林省农业科学院果树研究所 Pomology Research Institute，Jilin Academy of Agricultural Sciences	70.00	12.00	0.70	9月上中旬（吉林公主岭） Early to mid-September （Gongzhuling，Jilin）	施献举,1988
	象牙黄 Xiangya- huang	蛟河1号 × 金红 Jiaohe No.1 × Jinhong		120.00	13.00	0.94	7月下旬至8月上旬（吉林公主岭） From late July to early August （Gongzhuling，Jilin）	孙凤兰等,1998
	冰红 Binghong	金冠 × 红海棠 Golden Delicious × Red crabapple		64.00	12.40	0.49	9月上中旬（吉林公主岭） Early to mid-September （Gongzhuling，Jilin）	邵静等,2024
	塞外红 Saiwai- hong	不详 Parental origin unspecified	通辽市林业和草原研究所Tongliao Forestry and Grass land Science Research Institute	58.30	17.31	0.87	9月中下旬（内蒙古通辽） Mid-late September（Tongliao，Inner Mongolia）	王宝侠等,2014
	金萍 Jinping	宁酥 × 金红 Ningsu × Jinhong	辽宁农业职业技术学院 Liaoning Agricultural Vocational and Technical College	110.00	18.25	0.28	9月中旬（辽宁铁岭） Mid-September（Tieling，Liaoning）	高丹,2013
	紫香 Zixiang	赤阳 × 红海棠 Chiyang × Red crabapple	黑龙江省农业科学院浆果研究所 Berry Research Institute，Heilongjiang Academy of Agricultural Sciences	45.00	12.70	0.46	8月下旬（黑龙江宝清） Late August（Baoqing，Heilongjiang）	吴立仁等,2013
	龙丰 Longfeng	大秋 × 金红 Daqiu × Jinhong	黑龙江省农业科学院牡丹江分院 Mudanjiang Branch，Heilongjiang Academy of Agricultural Sciences	55.10	14.70	3.88	9月中旬（黑龙江牡丹江）Mid-September（Mudanjiang，Heilongjiang）	吴昆,1992
	龙红 Longhong	富士 × 铃铛 Fuji × Lingdang		75.00	16.10	0.37	8月下旬（黑龙江牡丹江） Late August （Mudanjiang，Heilongjiang）	刘延杰等,2002
	龙帅 Longshuai	金红 × 寒富 Jinhong × Hanfu		119.00	13.90	0.56	9月中旬（黑龙江牡丹江）Mid-September（Mudanjiang，Heilongjiang）	于文全等,2017
	龙冠 Longguan	七月鲜 × 金冠 Qiyuexian × Golden Delicious		94.50	13.00	0.17	9月上旬（黑龙江牡丹江） Early September（Mudanjiang，Heilongjiang）	李淑贤等,1987
	龙硕 Longshuo	龙丰自然芽变 Longfeng natural bud mutation		110.60	14.00	1.20	9月中下旬（黑龙江牡丹江）Mid-late September（Mudanjiang，Heilongjiang）	卜海东等,2023a
	龙艳 Longyan	金红自然芽变 Jinhong natural bud mutation		91.00	15.10	0.81	9月上中旬（黑龙江牡丹江）Early and middle September（Mudanjiang，Heilongjiang）	卜海东等,2023b
类型 Type	品种 Cultivar	亲本（父 × 母） Parent（male × female）	选育单位 Breeding organization	平均单果质量/g Average fruit weight	可溶性固形物/% Total soluble solid	总酸含量/% Total acid content	成熟期 Mature period	参考文献 Reference
小苹果 Crab apple	振兴1号 Zhenxing 1	龙冠 × 冻果 Longguan × Dongguo	黑龙江省农业科学院乡村振兴科技研究所 Institute of Rural Revitalization Technology，Heilongjiang Academy of Agricultural Sciences	43.00	16.50	0.48	9月下旬（黑龙江哈尔滨）Late September（Harbin，Heilongjiang）	焦奎宝等,2024

国家 Country	品种 Cultivar		亲本 Parent		选育单位 Breeding organization		特性 Characteristic	参考文献 Reference
英国 United kingdom	M26		M16 × M9		英国东茂林试验站 East Malling Research		半矮化，抗花叶病毒，抗寒Semi-dwarf，anti-mosaic virus，cold resistance	赵德英,2024a
英国 United kingdom	MM106		君袖 × M1 Junxiu × M1		英国约翰英尼斯园艺研究所和东茂林试验站John Innes Institute of Horticulture and East Malling Research		半矮化，抗苹果绵蚜，抗寒Semi-dwarf，resistant to apple aphid，cold resistance	赵德英,2024a
加拿大 Canada	SJM 44		Nertchinsk × M9		加拿大农业与农业食品园艺研究和发展中心 Agriculture and Agri-Food Canada，Atlantic Food and Horticulture Research Center		矮化，抗苹果绵蚜，抗寒 Dwarfing，resistance to apple aphid，cold resistance	赵德英,2024b
加拿大 Canada	SJP 84-5162		Robusta 5 × M27				矮化，抗白粉病、苹果绵蚜，抗寒Dwarfing，resistance to powdery mildew，apple aphid，cold resistance	赵德英,2024b
德国 Germany	Pillnitzer Supporter 4	M4 × M9		德国德累斯顿-皮尔尼茨植物遗传和作物研究 Leibniz Institute of Plant Genetics and Crop Plant Research，Dresden-Pieschen		半矮化、抗苹果绵蚜，较抗寒 Semi-dwarf，resistant to apple aphid，more resistant to cold		赵德英,2024c
俄罗斯 Russia	B9	M8 × Red Flag		俄罗斯米丘林斯克国立农业大学 Michurinsk State Agrarian University		矮化，抗颈腐病，抗寒Dwarfing，resistance to neck rot，cold resistance		赵德英,2024d
	71-3-150	58-257 × B9				半矮化，抗颈腐病，抗寒Semi-dwarf，neck rot resistance，cold resistance
	B118	Moscow Pear和M8与M9的混合花粉杂交Hybridization of Moscow Pear and M8 with mixed pollen of M9				半矮化，抗颈腐病和黑星病，极抗寒Semi-dwarf，resistant to neck rot and scab，extremely cold-resistant
	B10（62-396）	13-14 × B9				半矮化，抗火疫病，抗寒 Semi-dwarf，fire-resistant，cold-resistant
波兰 Poland	P22	M9 × Common Antonovka		波兰斯基尔涅维采果树研究所 Research Institute of Pomology and Floriculture，Skierniewice		极矮化，抗冠腐病和白粉病，极抗寒Extremely dwarf，resistant to crown rot and powdery mildew，and extremely cold-resistant.		赵德英,2024e
美国 USA	G935	Ottawa 3 × Robusta 5		美国康奈尔大学和Geneva试验站 Cornell University and Geneva Experiment Station		半矮化，高抗火疫病，抗寒Semi-dwarf，high fire resistance，cold resistance		赵德英,2024f
美国 USA	G814	Ottawa 3 × Robusta 5				半矮化，抗冠腐病，抗寒Semi-dwarf，crown rot resistance，cold resistance		赵德英,2024f
中国 China	CX4	M9 × 金红 M9 × Jinhong		中国农业科学院果树研究所Research Institute of Pomology，Chinese Academy of Agricultural Sciences		矮化，早果，抗寒 Dwarfing，early fruit，cold resistance		周家旭等,2024
	CX5	193 × M9				矮化，早果，抗寒 Dwarfing，early fruit，cold resistance		周家旭等,2024
	GM256	红海棠 × M9 Red crabapple × M9		吉林省农业科学院果树研究所 Pomology Research Institute，Jilin Academy of Agricultural Sciences		半矮化，早果，抗寒 Semi-dwarf，early fruit，cold resistance		张新宇等,1998
	GM310	红太平 × M9 Hongtaiping × M9				矮化，抗腐烂病，抗寒 Dwarfing，rot resistance，cold resistance		张冰冰等,2011
	ZM-2000	GM256自然实生 GM256 natural seeding				矮化，抗腐烂病，抗寒 Dwarfing，rot resistance，cold resistance		李粤渤等,2016
	辽砧2号 Liaozhen No.2	助列涅特 × M9 Zhulieniete × M9		辽宁省果树科学研究所 Liaoning Institute of Pomology		矮化，早果，易生根，抗寒Dwarfing，early fruit，easy to root，cold resistance		刘志等,2004
	77-34	小黄海棠 × M9 Small Yellow crabapple × M9		辽宁省果树科学研究所 Liaoning Institute of Pomology		半矮化，早果，抗寒 Semi-dwarf，early fruit，cold resistance		李恩生等,1990
	扎矮山荆子 Zhaai Malus baccata	山荆子自然实生 Natural seedling of Malus baccata		内蒙古自治区呼伦贝尔盟农业科学研究所Hulunbuir Agricultural Science Research Institute，Inner Mongolia Autonomous Region		矮化，抗腐烂病，抗寒 Dwarfing，rot resistance，cold resistance		孟庆炎等,1997
	SC6	SH5 × 黄海棠 SH5 × Yellow crabapple		山西农业大学园艺学院 College of Horticulture，Shanxi Agricultural University		矮化，早果，抗寒 Dwarfing，early fruit，cold resistance		蔚露等, 2024
	Y-1	晋西北野生山荆子实生选育 Seedling breeding of wild Malus baccata in northwest Shanxi		山西农业大学果树研究所 Pomology Institute，Shanxi Agricultural University		矮化，早果，抗旱，抗寒 Dwarfing，early fruit，drought resistance，cold resistance		周家旭等,2024
	Y-2					极矮化，早果，抗旱，抗寒 Extremely dwarf，early fruit，drought resistance，cold resistance
	MD001	山荆子 × M8 Malus baccata × M8		黑龙江省农业科学院 Heilongjiang Academy of Agricultural Sciences		半矮化，早果，抗寒 Semi-dwarf，early fruit，cold resistance
中国 China	KM矮化砧	花红 × M9 Huahong × M9		新疆生产建设兵团第七师农业科学研究所 Xinjiang Production and Construction Corps 7th Division Institute of Agricultural Sciences		矮化，早果，抗寒 Dwarfing，early fruit，cold resistance		李国萍等,2004
	SX126	M26 × 富平楸子 M26 × Fupingqiuzi		西北农林科技大学园艺学院 College of Horticulture，Northwest A & F University		矮化，抗旱，抗偏斜，抗寒 Dwarfing，drought resistance，deflection resistance，cold resistance		樊秀芳等,2005
	中砧1号 Zhongzhen 1	小金海棠自然实生 Small golden crabapple natural seedling		中国农业大学园艺学院 College of Horticulture，China Agricultural University		半矮化，抗落叶病，抗寒 Semi-dwarf，anti-defoliation，cold resistance		韩振海等,2013

国家 Country	品种 Cultivar		亲本 Parent		选育单位 Breeding organization		特性 Characteristic	参考文献 Reference
英国 United kingdom	M26		M16 × M9		英国东茂林试验站 East Malling Research		半矮化，抗花叶病毒，抗寒Semi-dwarf，anti-mosaic virus，cold resistance	赵德英,2024a
英国 United kingdom	MM106		君袖 × M1 Junxiu × M1		英国约翰英尼斯园艺研究所和东茂林试验站John Innes Institute of Horticulture and East Malling Research		半矮化，抗苹果绵蚜，抗寒Semi-dwarf，resistant to apple aphid，cold resistance	赵德英,2024a
加拿大 Canada	SJM 44		Nertchinsk × M9		加拿大农业与农业食品园艺研究和发展中心 Agriculture and Agri-Food Canada，Atlantic Food and Horticulture Research Center		矮化，抗苹果绵蚜，抗寒 Dwarfing，resistance to apple aphid，cold resistance	赵德英,2024b
加拿大 Canada	SJP 84-5162		Robusta 5 × M27				矮化，抗白粉病、苹果绵蚜，抗寒Dwarfing，resistance to powdery mildew，apple aphid，cold resistance	赵德英,2024b
德国 Germany	Pillnitzer Supporter 4	M4 × M9		德国德累斯顿-皮尔尼茨植物遗传和作物研究 Leibniz Institute of Plant Genetics and Crop Plant Research，Dresden-Pieschen		半矮化、抗苹果绵蚜，较抗寒 Semi-dwarf，resistant to apple aphid，more resistant to cold		赵德英,2024c
俄罗斯 Russia	B9	M8 × Red Flag		俄罗斯米丘林斯克国立农业大学 Michurinsk State Agrarian University		矮化，抗颈腐病，抗寒Dwarfing，resistance to neck rot，cold resistance		赵德英,2024d
	71-3-150	58-257 × B9				半矮化，抗颈腐病，抗寒Semi-dwarf，neck rot resistance，cold resistance
	B118	Moscow Pear和M8与M9的混合花粉杂交Hybridization of Moscow Pear and M8 with mixed pollen of M9				半矮化，抗颈腐病和黑星病，极抗寒Semi-dwarf，resistant to neck rot and scab，extremely cold-resistant
	B10（62-396）	13-14 × B9				半矮化，抗火疫病，抗寒 Semi-dwarf，fire-resistant，cold-resistant
波兰 Poland	P22	M9 × Common Antonovka		波兰斯基尔涅维采果树研究所 Research Institute of Pomology and Floriculture，Skierniewice		极矮化，抗冠腐病和白粉病，极抗寒Extremely dwarf，resistant to crown rot and powdery mildew，and extremely cold-resistant.		赵德英,2024e
美国 USA	G935	Ottawa 3 × Robusta 5		美国康奈尔大学和Geneva试验站 Cornell University and Geneva Experiment Station		半矮化，高抗火疫病，抗寒Semi-dwarf，high fire resistance，cold resistance		赵德英,2024f
美国 USA	G814	Ottawa 3 × Robusta 5				半矮化，抗冠腐病，抗寒Semi-dwarf，crown rot resistance，cold resistance		赵德英,2024f
中国 China	CX4	M9 × 金红 M9 × Jinhong		中国农业科学院果树研究所Research Institute of Pomology，Chinese Academy of Agricultural Sciences		矮化，早果，抗寒 Dwarfing，early fruit，cold resistance		周家旭等,2024
	CX5	193 × M9				矮化，早果，抗寒 Dwarfing，early fruit，cold resistance		周家旭等,2024
	GM256	红海棠 × M9 Red crabapple × M9		吉林省农业科学院果树研究所 Pomology Research Institute，Jilin Academy of Agricultural Sciences		半矮化，早果，抗寒 Semi-dwarf，early fruit，cold resistance		张新宇等,1998
	GM310	红太平 × M9 Hongtaiping × M9				矮化，抗腐烂病，抗寒 Dwarfing，rot resistance，cold resistance		张冰冰等,2011
	ZM-2000	GM256自然实生 GM256 natural seeding				矮化，抗腐烂病，抗寒 Dwarfing，rot resistance，cold resistance		李粤渤等,2016
	辽砧2号 Liaozhen No.2	助列涅特 × M9 Zhulieniete × M9		辽宁省果树科学研究所 Liaoning Institute of Pomology		矮化，早果，易生根，抗寒Dwarfing，early fruit，easy to root，cold resistance		刘志等,2004
	77-34	小黄海棠 × M9 Small Yellow crabapple × M9		辽宁省果树科学研究所 Liaoning Institute of Pomology		半矮化，早果，抗寒 Semi-dwarf，early fruit，cold resistance		李恩生等,1990
	扎矮山荆子 Zhaai Malus baccata	山荆子自然实生 Natural seedling of Malus baccata		内蒙古自治区呼伦贝尔盟农业科学研究所Hulunbuir Agricultural Science Research Institute，Inner Mongolia Autonomous Region		矮化，抗腐烂病，抗寒 Dwarfing，rot resistance，cold resistance		孟庆炎等,1997
	SC6	SH5 × 黄海棠 SH5 × Yellow crabapple		山西农业大学园艺学院 College of Horticulture，Shanxi Agricultural University		矮化，早果，抗寒 Dwarfing，early fruit，cold resistance		蔚露等, 2024
	Y-1	晋西北野生山荆子实生选育 Seedling breeding of wild Malus baccata in northwest Shanxi		山西农业大学果树研究所 Pomology Institute，Shanxi Agricultural University		矮化，早果，抗旱，抗寒 Dwarfing，early fruit，drought resistance，cold resistance		周家旭等,2024
	Y-2					极矮化，早果，抗旱，抗寒 Extremely dwarf，early fruit，drought resistance，cold resistance
	MD001	山荆子 × M8 Malus baccata × M8		黑龙江省农业科学院 Heilongjiang Academy of Agricultural Sciences		半矮化，早果，抗寒 Semi-dwarf，early fruit，cold resistance
中国 China	KM矮化砧	花红 × M9 Huahong × M9		新疆生产建设兵团第七师农业科学研究所 Xinjiang Production and Construction Corps 7th Division Institute of Agricultural Sciences		矮化，早果，抗寒 Dwarfing，early fruit，cold resistance		李国萍等,2004
	SX126	M26 × 富平楸子 M26 × Fupingqiuzi		西北农林科技大学园艺学院 College of Horticulture，Northwest A & F University		矮化，抗旱，抗偏斜，抗寒 Dwarfing，drought resistance，deflection resistance，cold resistance		樊秀芳等,2005
	中砧1号 Zhongzhen 1	小金海棠自然实生 Small golden crabapple natural seedling		中国农业大学园艺学院 College of Horticulture，China Agricultural University		半矮化，抗落叶病，抗寒 Semi-dwarf，anti-defoliation，cold resistance		韩振海等,2013

响应低温胁迫的类型 Types of response to low temperature stress	基因名称 Gene name	抗寒机制 Mechanism of cold resistance	参考文献 Reference
低温信号的感知与传导Low temperature signal perception and transduction	MdCPK1a	清除 ROS Scavenging of ROS	Dong et al.,2020
	MdbHLH4	促进MdCAX3L-2的表达，抑制低温诱导的Ca²⁺信号传导 Improve the expression of MdCAX3L-2 and inhibits Ca²⁺ signal transduction induced by low temperature	Yang et al.,2023
转录调控途径 Transcriptional regulatory pathway	MdMYB73	MdCBF2/3/4/5的表达水平增加，可溶性糖含量增加The expression level of MdCBF2/3/4/5 increased，and the soluble sugar content increased	Zhang et al.,2017
	MdbHLH33	正向调控MdCBF2表达，MdCOR15A-1和MdCOR15A-2的表达水平增加 Positive regulation of MdCBF2 expression increased the expression levels of MdCOR15A-1 and MdCOR15A-2	Xu et al.,2018
	MdCBF1/2	正向调控MdTST1/2表达，增加葡萄糖、果糖和蔗糖的含量 Positive regulation of MdTST1/2 expression enhances the accumulation of glucose，fructose，and sucrose	Li et al.,2024
	MdMYB88/124	正向调控MdCCA1激活MdCBF3的表达，促进花青素积累 Positive regulation of MdCCA1 activates MdCBF3 expression，thereby promoting anthocyanin accumulation	Xie et al.,2018
	MdNAC104	正向调控MdCBF1/3的表达 Positive regulation of MdCBF1/3 expression	Mei et al.,2023
	MdCBF3	正向调控MdGAD1表达，增加GABA合成 Positive regulation of MdGAD1 expression increased GABA synthesis	Liu et al.,2024a
	MdMYB23	正向调控MdANR，促进原花青素积累和ROS清除 Positive regulation of MdANR promotes proanthocyanidin accumulation and ROS scavenging	An et al.,2018
	MdBBX37	MdBBX37-MdICE1 协同激活MdCBF1表达 MdBBX37-MdICE1 synergistically activated MdCBF1 expression	An et al.,2021
	MdABI4	MdABI4与MdICE1相互作用，增强对MdCBF1的转录 MdABI4 interacts with MdICE1 to enhance the transcription of MdCBF1	An et al.,2022
转录调控途径 Transcriptional regulatory pathway	MdBIM1	正向调控MdCBF1表达，提高苹果抗寒性 Positive regulation of MdCBF1 expression promotes apple cold resistance	An et al.,2023
	MdCIbHLH1	MdCIbHLH1与MdERF1B相互作用，增强对MdCBF1的转录 MdCIbHLH1 interacts with MdERF1B to enhance the transcription of MdCBF1	Wang et al.,2021
	MdNAC104	促进MdCHS-b、MdCHI-a、MdF3Ha和MdANS-b表达，促进花青素积累 Promote the expression of MdCHS-b，MdCHI-a，MdF3H-a and MdANS-b，and promote anthocyanin accumulation	Mei et al.,2023
	MdWRKY40is	正向调控MdDFR，促进花青素积累 Positive regulation of MdDFR promotes anthocyanin accumulation	Xu et al.,2023d
	MdMRLK2	正向调控MdMYBPA1，促进MdANS和MdUFGT表达，诱导更多的花青素合成Positive regulation of MdMYBPA1 promotes the expression of MdANS and MdUFGT，thereby inducing enhanced anthocyanin biosynthesis	Jing et al.,2023a
	MdHY5	正向调控MdCBF1表达，提高苹果抗寒性；MdHY5与MdGH3-2/12和MdNCED2结合，降低苹果IAA/ABA Positive regulation of MdCBF1 expression promotes apple cold resistance；MdHY5 binds to MdGH3-2/12 and MdNCED2，and reduces the ratio of IAA/ABA in apple	An et al.,2017；Liu et al.,2024b
	MdARF17	POD和CAT活性升高 Increase POD and CAT activities	Shen et al.,2023a
	MdHYL1	正向调控MdMYB88/124表达，介导ROS清除、花青素和苯丙氨酸含量积累 Positive regulation of MdMYB88/124 expression mediates ROS scavenging，anthocyanin and phenylalanine accumulation	Shen et al.,2023b
	MdGolS5	增加肌醇半乳糖苷和棉子糖的含量，提高渗透水平 Increase the content of galactinol and raffinose，improve the penetration level	Xu et al.,2025
翻译后修饰调控途径 Post-translational modification regulation pathway	MdSIZ1	MdSIZ1对MdMYB1蛋白进行SUMO化修饰，提高MdMYB1蛋白的稳定性，促进花青苷的积累 MdSIZ1 SUMOylated MdMYB1 protein to improve the stability of MdMYB1 protein and promote the accumulation of anthocyanin	Zhou et al.,2017
	MdSIZ1	MdMYB2与MdSIZ1结合，促进对MdMYB1蛋白SUMO化修饰，促进花青苷的积累 MdMYB2 binds to MdSIZ1 to promote the SUMOylation of MdMYB1 protein and promote the accumulation of anthocyanins	Jiang et al.,2022
	MdPUB23	MdPUB23与MdICE1相互作用，促进MdICE1蛋白稳定性负调控冷应激反应 MdPUB23 interacts with MdICE1 to promote the stability of MdICE1 protein and negatively regulates cold stress response	Wang et al.,2022
	MdHDA6	MdHDA6对MdTCP15进行组蛋白去乙酰化并抑制其表达，提高苹果的抗寒性 MdHDA6 deacetylates histones of MdTCP15 and inhibits its expression to improve the cold resistance of apples	Guo et al.,2023