亚洲百合试管鳞茎膨大影响因子的研究

doi:10.16420/j.issn.0513-353x.2024-0368

摘要/Abstract

摘要： 以亚洲百合新品种‘甜心’和‘小海星’的组培小鳞茎为材料，研究了6-BA、NAA和草甘膦对鳞片不定芽诱导的效果，在此基础上分别研究了水杨酸（salicylic acid，SA），多效唑（paclobutrazol，PP₃₃₃）与2,4-表油菜素内酯（2,4-epibrassinolide，2,4-EBR）、蔗糖、活性炭（activated charcoal，AC）、NAA和6-BA以及基本培养基对小鳞茎膨大的影响，并分析小鳞茎在膨大发育过程中淀粉合成酶基因SSS和SSS2、糖外排转运蛋白基因SWEET14和SWEET15的表达变化。结果表明：‘甜心’和‘小海星’在MS + 0.5 mg · L^-16-BA + 0.2 mg · L^-1 NAA + 30 g · L^-1蔗糖培养基上出芽最多，分化系数分别为2.78、2.03。‘甜心’在MS + 0.1 mg · L^-1 SA+ 60 g · L^-1蔗糖、MS + 1.0 mg · L^-1 PP₃₃₃ + 0.05 mg · L^-1 2,4-EBR + 60 g · L^-1蔗糖和MS + 0.5 mg · L^-1 6-BA + 0.1 mg · L^-1 NAA + 60 g · L^-1蔗糖 + 1.0 g · L^-1AC培养基中的鳞茎膨大系数分别为4.42、5.15和4.16。筛选出其适宜的膨大培养基为MS + 1.0 mg · L^-1 NAA + 0.1 ~ 0.5 mg · L^-1 6-BA + 60 ~ 90 g · L^-1蔗糖 + 1.0 ~ 2.0 g · L^-1 AC。‘甜心’和‘小海星’在B5培养基下的膨大系数均显著高于MS和WPM培养基处理，分别为4.57和4.83。在B5培养基中SSS、SSS2和SWEET14、SWEET15随‘甜心’鳞茎的生长，表达量逐渐上升，在小鳞茎快速膨大期达到最高，叶片中在36 d即将进入小鳞茎快速膨大期时表达量最高。

关键词: 百合, 鳞茎膨大, 组织培养, 碳水化合物代谢

Abstract:

This study used tissue cultured bulblets of the new Asian lily cultivars‘Tianxin’and ‘Xiaohaixing’as materials to investigate the effects of 6-BA，NAA，and glyphosate on inducing adventitious buds on the scales. Based on the screening of suitable induction media for adventitious buds，the effects of plant growth regulators such as salicylic acid（SA），paclobutrazol（PP₃₃₃），and 2, 4-dichlorobrassinolide（2,4-EBR），sucrose，activated carbon（AC），NAA，6-BA，and basic media on the swelling of small bulbs were studied，and the expression changes of starch synthase genes SSS，SSS2，sugar efflux transporters genes SWEET14 and SWEET15 during the swelling development of small bulbs were analyzed. The results showed that‘Tianxin’and‘Xiaohaixing’germinated the most on MS + 0.5 mg · L^-1 6-BA + 0.2 mg · L^-1 NAA + 30 g · L^-1 sucrose medium，with differentiation coefficients of 2.78 and 2.03，respectively. The enlargement coefficients of‘Tianxin’in MS + 1.0 mg · L^-1 PP₃₃₃+ 0.05 mg · L^-1 2,4-EBR + 60 g · L^-1 sucrose，and MS + 0.5 mg · L^-1 6-BA + 0.1 mg · L^-1NAA + 60 g · L^-1 sucrose + 1.0 g · L^-1 activated carbon medium were 4.42，5.15，and 4.16，respectively. The suitable swelling medium for them was selected as MS + 1.0 mg · L^-1 NAA + 0.1-0.5 mg · L^-1 6-BA + 60-90 g · L^-1 sucrose + 1.0-2.0 g · L^-1 AC. The enlargement coefficients of‘Tianxin’and‘Xiaohaixing’in B5 medium were significantly higher than those in MS and WPM medium treatments，at 4.57 and 4.83，respectively. In B5 medium，the expression levels of soluble starch synthase genes SSS and SSS2，as well as sugar efflux transporters SWEET14 and SWEET15，gradually increase with the growth of the‘Tianxin’bulblet，reaching their highest levels during the rapid expansion phase of bulblet. In the leaves，the expression levels are highest at 36 days before entering the rapid expansion phase of bulblet.

Key words: Lilium, bulblet enlargement, tissue culture, carbohydrate metabolism

吴小丹, 孔祥凤, 高丽, 巩天耕, 朴美玲, 余鹏程, 贾桂霞. 亚洲百合试管鳞茎膨大影响因子的研究[J]. 园艺学报, 2025, 52(6): 1644-1660.

WU Xiaodan, KONG Xiangfeng, GAO Li, GONG Tiangeng, PIAO Meiling, YU Pengcheng, and JIA Guixia. Study on Influencing Factor of Bulblet Enlargement in vitro of Asiatic Lily[J]. Acta Horticulturae Sinica, 2025, 52(6): 1644-1660.

图/表 16

表1 用于RT-qPCR分析的引物序列

Table 1 Primers used for Real-time quantitative PCR analysis

基因名称 Gene name	正向引物（5′-3′） Forward primer sequence	反向引物（5′-3′） Reverse primer sequence
SSS（CL7226_C1）	AATGGTTGGAGGTGGAAATCT	GGCCTCCTTCGGAACATAAA
SSS2（CL3977_C2）	CACTGGTGGCCTTAGAGATAC	CCAGCATACTCTCCTTCGATAAA
SWEET14（CL255_C3）	TGACTCTTCTTGGTTGGATCTG	GGCATGAACTCGACACTCTT
SWEET15（CL7570_C1）	GGGTTCCAATCTCTTCCCTATG	TCCGATCGAGTTAATGGTGATG
Actin（内参基因 Reference gene）	CCCATTGAGCACGGCATTGTC	GGATTGAGAGGAGCTTCGGTGAGA

表2 NAA和6-BA对‘甜心’和‘小海星’百合鳞片诱导培养的影响

Table 2 Effects of NAA and 6-BA on scale-induced culture of‘Tianxin’and‘Xiaohaixing’lilies

处理 Treatment	生长调节剂/（mg · L^-1） Growth regulator			甜心 Tianxin			小海星 Xiaohaixing
处理 Treatment	6-BA	NAA	6-BA/ NAA	出芽时间/d Budding time	诱导率 Inductivity	分化系数 Coefficient of differentiation	出芽时间/d Budding time	诱导率 Inductivity	分化系数 Coefficient of differentiation
1	0.5	0.1	5.0	8 ~ 24	1.00 ± 0 a	2.44 ± 0.25 abc	13 ~ 30	0.90 ± 0.11 ab	1.92 ± 0.29 a
2	0.5	0.2	2.2	8 ~ 24	0.90 ± 0.09 ab	2.78 ± 0.69 a	13 ~ 30	0.89 ± 0.01 ab	2.03 ± 0.18 a
3	0.5	1.0	0.5	14 ~ 30	0.82 ± 0.08 abc	2.14 ± 0.29 abcd	24 ~ 45	0.37 ± 0.10 de	1.25 ± 0.45 bcd
4	1.0	0.1	10.0	14 ~ 30	0.96 ± 0.06 a	2.52 ± 0.63 ab	18 ~ 35	0.84 ± 0.05 abc	2.01 ± 0.16 a
5	1.0	0.2	5.0	14 ~ 30	0.89 ± 0.12 ab	2.12 ± 0.51 abcd	18 ~ 35	0.83 ± 0.13 abc	1.72 ± 0.54 ab
6	1.0	1.0	1.0	20 ~ 35	0.74 ± 0.18 bcd	1.55 ± 0.16 d	30 ~ 55	0.25 ± 0.15 ef	1.22 ± 0.40 bcd
7	1.5	0.1	15.0	18 ~ 35	0.92 ± 0.07 ab	2.73 ± 0.25 a	24 ~ 45	0.92 ± 0.06 a	1.81 ± 0.45 ab
8	1.5	0.2	7.5	18 ~ 35	0.89 ± 0.09 ab	2.09 ± 0.19 abcd	24 ~ 45	0.75 ± 0.13 bc	1.94 ± 0.65 a
9	1.5	1.0	1.5	24 ~ 45	0.63 ± 0.07 d	1.60 ± 0.40 d	30 ~ 50	0.19 ± 0.17 fg	1.01 ± 0.58 cd
10	2.0	0.1	20.0	24 ~ 45	0.83 ± 0.16 abc	2.46 ± 0.64 abc	24 ~ 45	0.71 ± 0.14 c	1.64 ± 0.38 abc
11	2.0	0.2	10.0	24 ~ 45	0.74 ± 0.12 bcd	1.82 ± 0.17 cd	24 ~ 45	0.47 ± 0.04 d	1.55 ± 0.41 abc
12	2.0	1.0	2.0	24 ~ 45	0.66 ± 0.22 cd	1.89 ± 0.32 bcd	40 ~ 65	0.08 ± 0.06 g	0.67 ± 0.52 d

表3 NAA和6-BA对‘甜心’和‘小海星’百合不定芽分化的方差分析

Table 3 Analysis of variance of NAA and 6-BA on adventitious bud differentiation of‘Tianxin’and‘Xiaohaixing’lilies

	因子Factor	平方和SS	自由度df	均方MS	F	P
甜心 Tianxin	6-BA	1.301	3	0.434	2.413	0.081
	NAA	4.832	2	2.416	13.443	0.000^**
	6-BA × NAA	1.774	6	0.296	1.645	0.161
	误差Error	7.009	39	0.180
	总计Total	254.532	51
小海星Xiaohaixing	6-BA	1.709	3	0.570	2.774	0.051
	NAA	8.709	2	4.354	21.204	0.000^**
	6-BA × NAA	0.456	6	0.076	0.370	0.894
	误差Error	9.857	48	0.205
	总计Total	158.494	60

表4 草甘膦对‘甜心’和‘小海星’百合鳞片诱导培养的影响

Table 4 Effect of glyphosate on the scale induction culture of‘Tianxin’and‘Xiaohaixing’lilies

草甘膦/mg · L^-1 Glyphosate	‘甜心’‘Tianxin’		‘小海星’‘Xiaohaixing’
草甘膦/mg · L^-1 Glyphosate	诱导率 Inductivity	分化系数 Coefficient of differentiation	诱导率 Inductivity	分化系数 Coefficient of differentiation
0（对照 Control）	0.89 ± 0 ab	2.26 ± 0.39 ab	0.89 ± 0.11 ab	2.59 ± 0.82
1.0	0.93 ± 0.13 a	2.70 ± 0.93 a	0.96 ± 0.06 a	3.56 ± 1.34
2.0	0.71 ± 0.06 bc	1.44 ± 0.12 b	0.78 ± 0.11 ab	2.51 ± 0.56
3.0	0.56 ± 0.12 c	1.30 ± 0.50 b	0.74 ± 0.06 b	2.33 ± 0.51

图1 不同浓度SA处理下‘甜心’和‘小海星’百合组培小鳞茎的生长

Fig. 1 The growth of‘Tianxin’and‘Xiaohaixing’lilies’s bulblet in vitro treated with different concentrations of SA

图2 不同SA浓度下‘甜心’和‘小海星’百合组培小鳞茎的膨大系数和新增小鳞茎个数不同小写字母表示同一品种不同处理间差异显著（P < 0.05）。下同

Fig. 2 The bulblet enlargement coefficient and the number of new bulblets added of‘Tianxin’and‘Xiaohaixing’lilies with different concentrations of SA in vitro Different lowercase letters of the same cultivar indicate significant differences among treatments（P < 0.05）. The same below

图3 不同浓度PP333及和2,4-EBR处理下‘甜心’和‘小海星’百合组培小鳞茎的生长

Fig. 3 The growth of‘Tianxin’and‘Xiaohaixing’lilies tissue culture bulblet treated with the different concentration of PP333 and 2,4-EBR

图4 不同浓度PP333及 2,4-EBR对‘甜心’和‘小海星’百合试管鳞茎膨大系数和新增小鳞茎个数的影响

Fig. 4 Effects of different concentrations of PP333 and 2,4-EBR on bulblet enlargement coefficient and the number of new bulblets added of ‘Tianxin’and‘Xiaohaixing’lilies in vitro

图5 ‘甜心’和‘小海星’组培小鳞茎在正交设计不同处理下小鳞茎的生长状况 1 ~ 9为试验组代号，详见表5

Fig. 5 The change of bulblet enlargement coefficient of tissue culture bulblet under Growth status of small bulblets 1-9 are the test group codes，as shown in table 5

表5 试管鳞茎膨大正交试验

Table 5 Orthogonal test of bulblet expansion in vitro

处理 Treatments	NAA/（mg · L^-1）	蔗糖/（g · L^-1） Sucrose	AC/（g · L^-1）	6-BA/（mg · L^-1）	膨大系数Expansion coefficient
处理 Treatments	NAA/（mg · L^-1）	蔗糖/（g · L^-1） Sucrose	AC/（g · L^-1）	6-BA/（mg · L^-1）	甜心Tianxin	小海星Xiaohaixing
1	0.1	30	0.5	0.1	2.67 ± 0.73 de	3.05 ± 1.26 de
2	0.1	60	1.0	0.5	4.16 ± 1.34 a	4.12 ± 1.47 abc
3	0.1	90	2.0	1.0	3.27 ± 1.54 bcd	3.85 ± 0.91 abcd
4	0.5	30	1.0	1.0	2.40 ± 0.60 e	2.95 ± 0.78 e
5	0.5	60	2.0	0.1	3.63 ± 1.24 abc	4.25 ± 1.28 abc
6	0.5	90	0.5	0.5	3.91 ± 1.64 ab	3.55 ± 0.69 bcde
7	1.0	30	2.0	0.5	2.96 ± 0.79 cde	4.35 ± 1.17 ab
8	1.0	60	0.5	1.0	2.79 ± 0.91 de	3.43 ± 0.88 cde
9	1.0	90	1.0	0.1	3.13 ± 1.11 bcde	4.43 ± 1.65 a

图6 不同基本培养基条件下‘甜心’和‘小海星’百合试管鳞茎膨大系数的变化不同小写字母表示不同的培养基处理间差异显著（P < 0.05）

Fig. 6 The changes of bulblet enlargement coefficient for‘Tianxin’and‘Xiaohaixing’lilies with different basic medium Different lowercase letters indicate significant differences between different culture medium treatments（P < 0.05）

图7 培养60 d‘甜心’和‘小海星’鳞茎膨大发育和叶片生长速度

Fig. 7 The bulblet enlargement and leaf growth rate of‘Tianxin’and‘Xiaohaixing’were cultured for 60 days

图8 ‘甜心’百合离体小鳞茎发育情况

Fig. 8 In vitro bulblet development of‘Tianxin’Lily

图9 鳞片横切和纵切细胞组织学观察 PC：薄壁细胞；EC：表皮细胞；VB：维管束组织；SG：淀粉粒

Fig. 9 Histological observation of squamosal transsection and slitting cells PC：Parenchyma cell；EC：Epidermal cells；VB：Vascular bundle tissue；SG：Starch granules

图10 ‘甜心’百合试管鳞片和叶片中SSS和SSS2在不同发育阶段的相对表达量不同小写字母表示相同基因不同的发育时间差异显著（P < 0.05），下同

Fig. 10 Relative expression levels of SSS and SSS2 in vitro scales and leaves of‘Tianxin’lilies at different developmental stages The different lowercase letters indicate significant differences between treatments at different developmental times of the same gene（P < 0.05），the same below

图11 ‘甜心’百合试管鳞片和叶片中SWEET14和SWEET15在不同发育阶段的相对表达量

Fig. 11 Relative expression levels of SWEET14 and SWEET15 of‘Tianxin’lilies in vitro scales and leaves at different developmental stages

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