枣77个品种果肉愈伤组织诱导培养研究

doi:10.16420/j.issn.0513-353x.2023-0205

摘要/Abstract

摘要：

果肉愈伤组织的大量培养可为悬浮细胞体系的建立、原生质体的分离与培养、遗传转化体系的建立及次生代谢产物的研究提供物质基础。以5个发育阶段的枣果肉为试材，共计77个枣品种，选用10种培养基进行枣果肉愈伤组织诱导研究。结果表明，基因型、果实发育阶段、培养基种类及其植物生长调节剂配比对枣果肉愈伤组织诱导效果均具有显著影响。在启动培养阶段，有42个品种分别筛选出1 ~ 6种可诱导出愈伤组织的培养基；有8种培养基可以成功诱导出愈伤组织，其中G1（4.41 g · L^-1 MS + 6 g · L^-1 琼脂 + 20 g · L^-1蔗糖 + 0.5 mg · L^-1 6-BA + 2.5 mg · L^-1 2,4-D）和G7（4.41 g · L^-1 MS + 6 g · L^-1琼脂 + 20 g · L^-1蔗糖 + 0.8 mg · L^-1 6-BA + 1.5 mg · L^-1 2,4-D）培养基的适用性最高，达60%，其次为G5（4.41 g · L^-1 MS + 6 g · L^-1琼脂 + 20 g · L^-1蔗糖 + 1.0 mg · L^-1 6-BA + 2.0 mg · L^-1 2,4-D），达45%；幼果最适于果肉愈伤组织培养，88%的品种可成功诱导出愈伤组织，随着果实成熟，果肉愈伤组织越不易诱导，在完全成熟的全红期只有‘圆铃小枣’成功诱导出愈伤组织。在继代培养阶段，有22个品种的愈伤组织实现了持续继代培养，适宜的培养基有G1（4.41 g · L^-1 MS + 6 g · L^-1琼脂 + 20 g · L^-1蔗糖 + 0.5 mg · L^-1 6-BA + 2.5 mg · L^-1 2,4-D）、G3（4.41 g · L^-1 MS + 6 g · L^-1 琼脂 + 20 g · L^-1蔗糖 + 1.0 mg · L^-1 6-BA + 1.0 mg · L^-1 2,4-D）、G5（4.41 g · L^-1 MS + 6 g · L^-1 琼脂 + 20 g · L^-1 蔗糖 + 1.0 mg · L^-1 6-BA + 2.0 mg · L^-1 2,4-D）、G6（4.41 g · L^-1 MS + 6 g · L^-1 琼脂 + 20 g · L^-1 蔗糖 + 3.0 mg · L^-1 TDZ + 0.5 mg · L^-1 NAA）、G7（4.41 g · L^-1 MS + 6 g · L^-1琼脂 + 20 g · L^-1蔗糖 + 0.8 mg · L^-1 6-BA + 1.5 mg · L^-1 2,4-D）、G9（4.41 g · L^-1 MS + 6 g · L^-1琼脂 + 20 g · L^-1蔗糖 + 0.5 mg · L^-1 NAA + 1.0 mg · L^-1 6-BA）和G10（4.41 g · L^-1 MS + 6 g · L^-1琼脂 + 25 g · L^-1蔗糖 + 0.4 mg · L^-1 6-BA + 1.5 mg · L^-1 2,4-D）；改变植物生长调节剂配比可以显著提高愈伤继代效果。此外，在启动培养阶段，有极少量的品种，如‘无核72’和‘临猗梨枣’可以直接诱导出芽。

关键词: 枣, 愈伤组织, 果肉培养, 植物生长调节剂

Abstract:

Mass culture of pulp callus can provide material basis for the establishment of suspension cell system，the isolation and culture of protoplasts，the establishment of genetic transformation system and the study of secondary metabolites. In this study，10 media were selected to study callus induction of jujube pulp from 77 jujube cultivars at five developmental stages. The results showed that the genotypes，fruit development stages，medium type and the proportion of growth regulators had significant effects on the callus induction. At the initial cultivating stage，one to six kinds of medium were screened which were suitable for callus induction for 42 cultivars. There were eight kinds of media that can successfully induce callus，among which G1（4.41 g · L^-1 MS + 6 g · L^-1 agar + 20 g · L^-1 sucrose + 0.5 mg · L^-1 6-BA + 2.5 mg · L^-1 2,4-D）and G7（4.41 g · L^-1 MS + 6 g · L^-1 agar + 20 g · L^-1 sucrose + 0.8 mg · L^-1 6-BA + 1.5 mg · L^-1 2,4-D）medium had the highest applicability，up to 60%，followed by G5（4.41 g · L^-1 MS + 6 g · L^-1 agar + 20 g · L^-1 sucrose + 1.0 mg · L^-1 6-BA + 2.0 mg · L^-1 2,4-D），up to 45%. Young fruits were the most suitable for pulp callus induction and cultivation，and the successful rate of cultivars reached 88%. In addition，the closer the fruit was to maturity，the more difficult the pulp callus cultivation was. Among them，only‘Yuanling Xiaozao’pulp at the whole red stage could be successfully induced callus. At the sub-cultivation stage，22 cultivars pulp callus were achieved subcultivition，and the appropriate medium was G1（4.41 g · L^-1 MS + 6 g · L^-1 agar + 20 g · L^-1 sucrose + 0.5 mg · L^-1 6-BA + 2.5 mg · L^-1 2,4-D），G3（4.41 g · L^-1 MS + 6 g · L^-1 agar + 20 g · L^-1 sucrose + 1.0 mg · L^-1 6-BA + 1.0 mg · L^-1 2,4-D），G5（4.41 g · L^-1 MS + 6 g · L^-1 agar + 20 g · L^-1 sucrose + 1.0 mg · L^-1 6-BA + 2.0 mg · L^-1 2,4-D），G6（4.41 g · L^-1 MS + 6 g · L^-1 agar + 20 g · L^-1 sucrose + 3.0 mg · L^-1 TDZ + 0.5 mg · L^-1 NAA），G7（4.41 g · L^-1MS + 6 g · L^-1 agar + 20 g · L^-1 sucrose + 0.8 mg · L^-1 6-BA + 1.5 mg · L^-1 2,4-D），G9（4.41 g · L^-1 MS + 6 g · L^-1 agar + 20 g · L^-1 sucrose + 0.5 mg · L^-1 NAA + 1.0 mg · L^-16-BA），G10（4.41 g · L^-1 MS + 6 g · L^-1 agar + 25 g · L^-1 sucrose + 0.4 mg · L^-1 6-BA + 1.5 mg · L^-1 2,4-D）. The changes of hormone ratio could significantly improve the callus sub-cultivation effect at the following cultivating stage. In addition，at the initial cultivating，a very small number of cultivars such as‘Wuhe 72’and‘Linyi Lizao’could directly induce bud germination.

Key words: Chinese jujube, callus, pulp cultivation, plant growth regulator

赵若雨, 郭晓雪, 朱文慧, 李瑞梅, 赵佳雪, 杜丹, 代丽, 刘志国, 王立新, 刘孟军. 枣77个品种果肉愈伤组织诱导培养研究[J]. 园艺学报, 2024, 51(4): 903-914.

ZHAO Ruoyu, GUO Xiaoxue, ZHU Wenhui, LI Ruimei, ZHAO Jiaxue, DU Dan, DAI Li, LIU Zhiguo, WANG Lixin, LIU Mengjun. Induction and Cultivation of 77 Jujube Cultivars Pulp Callus[J]. Acta Horticulturae Sinica, 2024, 51(4): 903-914.

图/表 12

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品种编号及名称 Number and cultivars	培养基编号 Medium No.					合计 Total
品种编号及名称 Number and cultivars	幼果期 Young fruit stage	白熟期 White stage	点红期 Dot red stage	半红期 Half red stage	全红期 Full red stage	合计 Total
1临泽大枣 Linze Dazao	G1^*，G6，G10	—	—	—	—	3
2鸭枣 Yazao	G1，G3^*，G5，G10	—	—	—	—	4
3婆婆枣 Popozao	G3，G5，G6，G9^*	G3	—	—	—	4
4壶瓶枣 Hupingzao	G1，G5，G6，G7^*，G10	G7	G7	—	—	5
5圆铃小枣 Yuanling Xiaozao	G1，G3，G6，G7，G10^*	—	—	G3	G3	5
6朝阳圆枣 Chaoyang Yuanzao	G1，G7^*，G10	G7	—	—	—	3
7三棱枣 Sanlengzao	G1，G6，G7^*	—	G7	—	—	3
8无核枣76 Wuhe 76	G1，G7^*，G10	G7	—	—	—	3
9小紫枣 Xiaozizao	G1，G3，G5，G6^*，G7，G10	G1	—	G1，G5，G6	—	6
10绵小枣 Mianxiaozao	G1^*，G3，G5，G10	G1	G3，G9	—	—	5
11临汾蜜枣 Linfen Mizao	G1^*，G5，G6	G6	—	—	—	3
12酸枣 Suanzao	G1^*，G9	—	—	—	—	2
13晚棉枣 Wanmianzao	G3，G5，G7，G9，G10^*	G5	—	—	—	5
14茶壶枣 Chahuzao	G7^*，G9，G10	G7	—	—	—	3
15相枣 Xiangzao	G3^*，G5，G7，G9，G10	G3	—	—	—	5
16连县木枣 Lianxian Muzao	G1，G10^*	—	—	—	—	2
17扁核酸 Bianhesuan	G9	—	—	—	—	1
18鸡蛋枣 Jidanzao	G1，G7，G9^*	—	—	—	—	3
19不落酥 Buluosu	G1，G3，G7^*，G10	G10	—	—	—	4
20稷山板枣 Jishan Banzao	G1，G3，G5，G7，G9，G10^*	G1，G10	—	G3，G10	—	6
21馒馒枣 Manmanzao	G10	—	—	—	—	1
22大叶无核 Daye Wuhe	G1^*，G3，G6	G6	—	G3	—	3
23宣城圆枣 Xuancheng Yuanzao	G3^*，G5，G6，G7	G7	—	—	—	4
24无核72 Wuhe 72	G1，G5，G7^*，G9	—	—	—	—	4
25阜香 Fuxiang	G6	—	—	—	—	1
26早脆王 Zaocuiwang	G5^*，G7	—	—	—	—	2
27颖秀 Yingxiu	G5，G6^*	—	—	—	—	2
28饼饼罐 Bingbingguan	G5，G7^*，G10	—	—	—	—	3
29牛角枣 Niujiaozao	G1^*，G7，G9	—	—	—	—	3
30金丝小枣 Jinsi Xiaozao	G1，G7，G10^*	—	—	—	—	3
31狗头枣 Goutouzao	G1，G5^*，G6，G7，G9，G10	—	—	—	—	6
32紫圆枣 Ziyuanzao	G1^*，G5，G6	—	—	—	—	3
33紫枣 Zizao	G5	—	—	—	—	1
34临猗梨枣 Linyi Lizao	G1，G9^*	—	—	—	—	2
35大荔龙枣 Dali Longzao	G5^*，G6，G7	—	—	G5	—	3
36星光 Xingguang	G3，G7，G10^*	G3	—	—	—	3
37南京大木 Nanjing Damu	G7，G9^*	—	—	—	—	2
38赞新大枣 Zanxin Dazao	G1，G7，G10^*	—	—	—	—	3
39傻枣 Shazao	G5	—	—	—	—	1
40无核丰 Wuhefeng	G3	—	—	G3	—	1
41酸枣2X Suanzao 2X	G3，G7^*	G3，G7	—	G3	—	2
42无核68 Wuhe 68	G9	—	—	—	—	1

品种编号及名称 Number and cultivars	培养基编号 Medium No.					合计 Total
品种编号及名称 Number and cultivars	幼果期 Young fruit stage	白熟期 White stage	点红期 Dot red stage	半红期 Half red stage	全红期 Full red stage	合计 Total
1临泽大枣 Linze Dazao	G1^*，G6，G10	—	—	—	—	3
2鸭枣 Yazao	G1，G3^*，G5，G10	—	—	—	—	4
3婆婆枣 Popozao	G3，G5，G6，G9^*	G3	—	—	—	4
4壶瓶枣 Hupingzao	G1，G5，G6，G7^*，G10	G7	G7	—	—	5
5圆铃小枣 Yuanling Xiaozao	G1，G3，G6，G7，G10^*	—	—	G3	G3	5
6朝阳圆枣 Chaoyang Yuanzao	G1，G7^*，G10	G7	—	—	—	3
7三棱枣 Sanlengzao	G1，G6，G7^*	—	G7	—	—	3
8无核枣76 Wuhe 76	G1，G7^*，G10	G7	—	—	—	3
9小紫枣 Xiaozizao	G1，G3，G5，G6^*，G7，G10	G1	—	G1，G5，G6	—	6
10绵小枣 Mianxiaozao	G1^*，G3，G5，G10	G1	G3，G9	—	—	5
11临汾蜜枣 Linfen Mizao	G1^*，G5，G6	G6	—	—	—	3
12酸枣 Suanzao	G1^*，G9	—	—	—	—	2
13晚棉枣 Wanmianzao	G3，G5，G7，G9，G10^*	G5	—	—	—	5
14茶壶枣 Chahuzao	G7^*，G9，G10	G7	—	—	—	3
15相枣 Xiangzao	G3^*，G5，G7，G9，G10	G3	—	—	—	5
16连县木枣 Lianxian Muzao	G1，G10^*	—	—	—	—	2
17扁核酸 Bianhesuan	G9	—	—	—	—	1
18鸡蛋枣 Jidanzao	G1，G7，G9^*	—	—	—	—	3
19不落酥 Buluosu	G1，G3，G7^*，G10	G10	—	—	—	4
20稷山板枣 Jishan Banzao	G1，G3，G5，G7，G9，G10^*	G1，G10	—	G3，G10	—	6
21馒馒枣 Manmanzao	G10	—	—	—	—	1
22大叶无核 Daye Wuhe	G1^*，G3，G6	G6	—	G3	—	3
23宣城圆枣 Xuancheng Yuanzao	G3^*，G5，G6，G7	G7	—	—	—	4
24无核72 Wuhe 72	G1，G5，G7^*，G9	—	—	—	—	4
25阜香 Fuxiang	G6	—	—	—	—	1
26早脆王 Zaocuiwang	G5^*，G7	—	—	—	—	2
27颖秀 Yingxiu	G5，G6^*	—	—	—	—	2
28饼饼罐 Bingbingguan	G5，G7^*，G10	—	—	—	—	3
29牛角枣 Niujiaozao	G1^*，G7，G9	—	—	—	—	3
30金丝小枣 Jinsi Xiaozao	G1，G7，G10^*	—	—	—	—	3
31狗头枣 Goutouzao	G1，G5^*，G6，G7，G9，G10	—	—	—	—	6
32紫圆枣 Ziyuanzao	G1^*，G5，G6	—	—	—	—	3
33紫枣 Zizao	G5	—	—	—	—	1
34临猗梨枣 Linyi Lizao	G1，G9^*	—	—	—	—	2
35大荔龙枣 Dali Longzao	G5^*，G6，G7	—	—	G5	—	3
36星光 Xingguang	G3，G7，G10^*	G3	—	—	—	3
37南京大木 Nanjing Damu	G7，G9^*	—	—	—	—	2
38赞新大枣 Zanxin Dazao	G1，G7，G10^*	—	—	—	—	3
39傻枣 Shazao	G5	—	—	—	—	1
40无核丰 Wuhefeng	G3	—	—	G3	—	1
41酸枣2X Suanzao 2X	G3，G7^*	G3，G7	—	G3	—	2
42无核68 Wuhe 68	G9	—	—	—	—	1

级别 Grade	状态 Status	颜色 Colour	增殖能力 Proliferating ability	果肉愈伤组织可继续增殖生长的枣品种 Cultivars whose pulp callus can continue to proliferate and grow
1	松软 Soft	乳白色，黄白色 Milk-white， yellow-white	较强 Stronger	颖秀，阜香，三棱枣、小紫枣，无核76，不落酥，饼饼罐，临汾蜜枣 Yingxiu，Fuxiang，Sanlengzao，Xiaozizao，Wuhe 76，Buluosu，Bingbingugan，Linfen Mizao
2	松软 Soft	黄白色 Yellow-white	强 Strong	相枣，绵小枣，晚棉枣（半红期），大叶无核，小紫枣（半红期），绵小枣（点红期） Xiangzao，Mianxiaozao，Wanmianzao（Half red stage），Daye Wuhe，Xiaozizao（Half red stage），Mianxiaozao（Dot red stage）
3	松软 Soft	深黄色 Deep yellow	一般 Ordinary	圆铃小枣，早脆王，茶壶枣，无核72，酸枣2X，壶瓶枣，稷山板枣，稷山板枣（半红期） Yuanling Xiaozao，Zaocuiwang，Chahuzao，Wuhe 72，Suanzao 2X，Hupingzao，Jishan Banzao，Jishan Banzao（Half red stage）
4	紧密 Tight	黄褐色 Tawny	较弱 Weaker	宣城圆枣，酸枣，金丝小枣，无核丰（半红期）、酸枣2X（半红期） Xuancheng Yuanzao，Suanzao，Jinsi Xiaozao，Wuhefeng（Half red stage），Suanzao 2X（Half red stage）
5	紧密 Tight	黑褐色 Black brown	弱 Weak	鸭枣，朝阳圆枣，馒馒枣，星光（白熟期），圆铃小枣（全红期） Yazao，Chaoyang Yuanzao，Manmanzao，Xingguang（White stage）， Yuanling Xiaozao（Full red stage）

级别 Grade	状态 Status	颜色 Colour	增殖能力 Proliferating ability	果肉愈伤组织可继续增殖生长的枣品种 Cultivars whose pulp callus can continue to proliferate and grow
1	松软 Soft	乳白色，黄白色 Milk-white， yellow-white	较强 Stronger	颖秀，阜香，三棱枣、小紫枣，无核76，不落酥，饼饼罐，临汾蜜枣 Yingxiu，Fuxiang，Sanlengzao，Xiaozizao，Wuhe 76，Buluosu，Bingbingugan，Linfen Mizao
2	松软 Soft	黄白色 Yellow-white	强 Strong	相枣，绵小枣，晚棉枣（半红期），大叶无核，小紫枣（半红期），绵小枣（点红期） Xiangzao，Mianxiaozao，Wanmianzao（Half red stage），Daye Wuhe，Xiaozizao（Half red stage），Mianxiaozao（Dot red stage）
3	松软 Soft	深黄色 Deep yellow	一般 Ordinary	圆铃小枣，早脆王，茶壶枣，无核72，酸枣2X，壶瓶枣，稷山板枣，稷山板枣（半红期） Yuanling Xiaozao，Zaocuiwang，Chahuzao，Wuhe 72，Suanzao 2X，Hupingzao，Jishan Banzao，Jishan Banzao（Half red stage）
4	紧密 Tight	黄褐色 Tawny	较弱 Weaker	宣城圆枣，酸枣，金丝小枣，无核丰（半红期）、酸枣2X（半红期） Xuancheng Yuanzao，Suanzao，Jinsi Xiaozao，Wuhefeng（Half red stage），Suanzao 2X（Half red stage）
5	紧密 Tight	黑褐色 Black brown	弱 Weak	鸭枣，朝阳圆枣，馒馒枣，星光（白熟期），圆铃小枣（全红期） Yazao，Chaoyang Yuanzao，Manmanzao，Xingguang（White stage）， Yuanling Xiaozao（Full red stage）

发育期 Developmental stages	供试品种总数 Total number of tested cultivars	诱导出愈伤组织的品种数 Number of cultivars that induced callus	诱导成功率/% Successful induction rate
幼果期 Young fruit stage	48	42	87.50
白熟期 White stage	36	16	44.44
点红期 Dot red stage	37	3	8.11
半红期 Half red stage	46	8	17.39
全红期 Full red stage	64	1	1.56