不同种源的葡萄种子休眠及其解除的研究

doi:10.3724/SP.J.1003.2008.08049

摘要/Abstract

摘要：

为探讨葡萄种子休眠与解除的规律, 我们选择起源于东亚、北美-中美、欧洲-中亚3个分布中心的不同种类及其杂种的20个品种, 研究了它们成熟种子的外部形态与萌发行为, 种子的休眠特性与休眠解除的方法, 并模拟四季温度的交替变化研究了环境温度对种子休眠的影响。结果表明, 不同起源的各类葡萄种子的休眠类型均为生理休眠, 但其休眠程度不同, 休眠解除方式也存在差异。其中欧亚种和东亚种的种子休眠较浅, 美洲种种子休眠较深; 杂交种比亲本所属类别的种子休眠程度浅。对于欧亚种、东亚种及其杂交种(欧山杂种)而言, 5ºC冷层积和变温层积(即20ºC (14 h) /10ºC (10 h)和30ºC (14 h) /20ºC (10 h))2个月能够有效地或部分解除它们的种子休眠; 但对美洲种和欧美杂种而言, 仅5ºC冷层积且层积时间需要延长至6个月才能解除其休眠, 变温层积和25ºC暖层积都不能解除休眠。四季温度的交替变化模拟实验进一步证明了不同起源的葡萄种子的休眠程度不同。这些休眠特性及其解除方式反映了不同起源葡萄种类的环境适应性。本文研究结果为葡萄资源的引种和育种提供了参考数据。

关键词: 葡萄种子, 休眠, 遗传背景

Abstract

To investigate dormancy and release mechanisms of grape seeds, we compared seed morphological characteristics and germination behavior of different grape varieties and their dormancy release under different conditions including an alternative temperature change simulation for a season cycle. The grape varieties tested included species introduced from three distribution centers (America, Europe-Asia and East-Asia) and their cross breeds. Seed dormancy of all varieties was attributed to physiological dormancy, but varied in degree of dormancy. The dormancy-releasing requirements of different varieties also differed, correlating to their genetic backgrounds. As a whole, seeds of Europe-Asia species and East-Asia species exhibited a lower degree of dormancy, compared to a relatively high degree of dormancy for American species. For Europe-Asia species, East-Asia species and their cross breeds, seed dormancy was effectively broken by cold stratification (5ºC) and partially broken by alternative temperature stratificationviz. 20ºC (14 h) /10ºC (10 h) and 30ºC (14 h) /20ºC (10 h) for two months. As for American species and cross breeds with Europe-Asia species, seed dormancy could only be broken by more than six-month cold stratification; neither alternative temperature stratification nor warm stratification (25ºC) affected seed dormancy release. Experiments simulating seasonal alternative changes further showed that seed dormancy degrees were related to their proveniences and adaptations to environment. Our results provide basic data for germplasm introduction and breeding of grapes.

Key words: Vitis seed, dormancy, genetic background

甘阳英, 李绍华, 宋松泉, 王伟青, 程红焱 (2008) 不同种源的葡萄种子休眠及其解除的研究. 生物多样性, 16, 570-577. DOI: 10.3724/SP.J.1003.2008.08049.

Yangying Gan, Shaohua Li, Songquan Song, Weiqing Wang, Hongyan Cheng (2008) Seed dormancy and release of grapes from different proveniences. Biodiversity Science, 16, 570-577. DOI: 10.3724/SP.J.1003.2008.08049.

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链接本文: https://www.biodiversity-science.net/CN/10.3724/SP.J.1003.2008.08049

https://www.biodiversity-science.net/CN/Y2008/V16/I6/570

图/表 8

表1 实验材料的描述

Table 1 Description of the experimental materials

种类 Category	选用品种 Variety	特性 Remarks
东亚种 (S) East-Asia species	'通化一号' Tonghuayi '双优' Shuangyou '左山一' Zuoshanyi '左山二' Zuoshaner '双红' Shuanghong '双丰' Shuangfeng	均为中国原生种山葡萄的选育品种, 产于中国东北、华北及华东部分地区, 分布海拔为200-2,100 m, 在葡萄属中抗寒性最强 (耐受-40℃) All of these varieties are selected from wild grape of V. amurensis, a Chinese origin species, distributed at altitudes of 200-2,100 m in northeast of China, northern China and parts of eastern China, the most cold-resistant species of grape, able to survive at -40℃.
美洲种 (M) American species	'康可' Concord '贝达' Beta	美洲葡萄实生苗选育品种 Selected from seedlings of V. labrusca, a wild American species 美洲种杂种 A cross-breed within American species (V. vulpine×V. labrusca)
欧亚种 (O) Europe-Asia species	'京秀' Jingxiu '京玉' Jingyu '京蜜' Jingmi '玫瑰香' Muscat Hamburg '96-10' '意大利' Italia	均为欧洲葡萄的选育品种。'京秀'、'京玉'、'京蜜'为早熟品种, '玫瑰香'为中熟品种, '96-10'、'意大利'为晚熟品种。 All of these varieties are selected from V.vinifera, the only existing species of Europe-Asia origin. Jingxiu, Jingyu, and Jingmi are early-maturing. Muscat Hamburg is medium-maturing, and 96-10 and Italia are late-maturing.
欧山杂种 (O×S) Cross-breed of Europe-Asia species × East-Asia species	'北醇' Beichun '北红' Beihong	欧洲葡萄与东亚种山葡萄的杂交后代 Cross-breed of Europe-Asia species × East-Asia species
欧美杂种 (O×M) Cross-breed of Europe-Asia species × American species	'京亚' Jingya '高妻' Takqtsuma	欧洲葡萄和美洲葡萄的杂交后代 Cross-breed of Europe-Asia species × American species
蘡欧杂种 (Y×O) Cross-breed of East-Asia species×Europe-Asia species	'北紫' Beizi '北丰' Beifeng	东亚种蘡薁葡萄和欧洲葡萄的杂交后代。蘡薁葡萄 (V. thunbergii) 为中国原生种, 产于华北、华中、华东、西南和华南地区, 分布于海拔150-2,500 m, 是中国分布最广的葡萄种。 Cross-breed of East-Asia species×Europe-Asia species.V. thunbergiiis a wild grape species of Chinese origin, most widely distributed in China. It distributes at altitudes of 150-2,500 m in northern China, central China, eastern China, southwest of China, and southern China.

表1 实验材料的描述

Table 1 Description of the experimental materials

种类 Category	选用品种 Variety	特性 Remarks
东亚种 (S) East-Asia species	'通化一号' Tonghuayi '双优' Shuangyou '左山一' Zuoshanyi '左山二' Zuoshaner '双红' Shuanghong '双丰' Shuangfeng	均为中国原生种山葡萄的选育品种, 产于中国东北、华北及华东部分地区, 分布海拔为200-2,100 m, 在葡萄属中抗寒性最强 (耐受-40℃) All of these varieties are selected from wild grape of V. amurensis, a Chinese origin species, distributed at altitudes of 200-2,100 m in northeast of China, northern China and parts of eastern China, the most cold-resistant species of grape, able to survive at -40℃.
美洲种 (M) American species	'康可' Concord '贝达' Beta	美洲葡萄实生苗选育品种 Selected from seedlings of V. labrusca, a wild American species 美洲种杂种 A cross-breed within American species (V. vulpine×V. labrusca)
欧亚种 (O) Europe-Asia species	'京秀' Jingxiu '京玉' Jingyu '京蜜' Jingmi '玫瑰香' Muscat Hamburg '96-10' '意大利' Italia	均为欧洲葡萄的选育品种。'京秀'、'京玉'、'京蜜'为早熟品种, '玫瑰香'为中熟品种, '96-10'、'意大利'为晚熟品种。 All of these varieties are selected from V.vinifera, the only existing species of Europe-Asia origin. Jingxiu, Jingyu, and Jingmi are early-maturing. Muscat Hamburg is medium-maturing, and 96-10 and Italia are late-maturing.
欧山杂种 (O×S) Cross-breed of Europe-Asia species × East-Asia species	'北醇' Beichun '北红' Beihong	欧洲葡萄与东亚种山葡萄的杂交后代 Cross-breed of Europe-Asia species × East-Asia species
欧美杂种 (O×M) Cross-breed of Europe-Asia species × American species	'京亚' Jingya '高妻' Takqtsuma	欧洲葡萄和美洲葡萄的杂交后代 Cross-breed of Europe-Asia species × American species
蘡欧杂种 (Y×O) Cross-breed of East-Asia species×Europe-Asia species	'北紫' Beizi '北丰' Beifeng	东亚种蘡薁葡萄和欧洲葡萄的杂交后代。蘡薁葡萄 (V. thunbergii) 为中国原生种, 产于华北、华中、华东、西南和华南地区, 分布于海拔150-2,500 m, 是中国分布最广的葡萄种。 Cross-breed of East-Asia species×Europe-Asia species.V. thunbergiiis a wild grape species of Chinese origin, most widely distributed in China. It distributes at altitudes of 150-2,500 m in northern China, central China, eastern China, southwest of China, and southern China.

表2 不同种源的葡萄品种成熟种子的千粒重和最初萌发率

Table 2 1000-seed weight and initial germination percentage of mature seeds of grape varieties from different proveniences

种类 Category	品种 Variety	千粒重 1000-seed weight (g)	最初萌发率 Initial germination (%)
东亚种 (S)	'双红' Shuanghong	27.6^d	0
	'通化一号' Tonghuayi	33.7^c	0
	'双优' Shuangyou	27.3^d	0
	'左山一' Zuoshanyi	36.7^c	0
	'左山二' Zuoshaner	28.9^d	0
	'双丰' Shuangfeng	28.7^d	0
美洲种 (M)	'康可' Concord	52.3^b	0
美洲种 (M)	'贝达' Beta	38.6^c	0
欧亚种 (O)	'京秀' Jingxiu	56.8^b	46.3±6.0^a
	'京玉' Jingyu	32.9^d	0
	'玫瑰香' Muscat Hamburg	40.9^c	0
	'96-10'	57.9^b	0
	'意大利' Italia	41.5^c	0
欧山杂种 (O×S)	'北醇' Beichun	41.9^c	16.7±2.6^b
欧山杂种 (O×S)	'北红' Beihong	42.3^c	14.3±1.6^b
欧美杂种 (O×M)	'京亚' Jingya	90.0^a	0
欧美杂种 (O×M)	'高妻' Takqtsuma	103.8^a	0
蘡欧杂种 (Y×O)	'北紫' Beizi	34.9^c	16±4^b
蘡欧杂种 (Y×O)	'北丰' Beifeng	38.4^c	20±6.9^b

图1 不同种源的葡萄品种成熟种子和种胚的外观形态。A: 种子背面观(上排)和种子腹面观(下排); B: 种胚。

Fig. 1 Morphological appearance of mature seeds of grape varieties from different proveniences. A, Hilum side (above line) and back side (below line) of seeds; B, Embryos.

图2 不同种源葡萄品种的种子在吸胀过程中含水量(g H2O g-1 DW)的变化。A: 山葡萄品种; B: 北醇(欧山杂种)、京秀和京玉(欧亚种)、京亚(欧美杂种)、康可(美洲种)。

Fig. 2 Changes in seed water contents (g H2O g-1 DW) during imbibition of grape varieties from different proveniences. A, Varieties of East-Asia species (S); B, Beichun (O×S), Jingxiu and Jingyu (O), Jingya (O×M) and Concord (M).

图3 浓硫酸处理对葡萄种子萌发率的影响。贝达(美洲种); 北红(欧山杂种); 左山一(山葡萄); 京玉(欧亚种); 京亚(欧美杂种)。

Fig. 3 Effects of sulfuric acid treatment on germination of grape seeds. Beta (M); Beihong (O×S); Zuoshanyi (S); Jingyu (O); Jingya (O×M).

图4 除去种胚端种皮对种子萌发的影响和离体胚的萌发。A, 除去种胚端种皮的种子在潮湿的滤纸上萌发; B, 离体胚在MS培养基上萌发(不含任何外源植物激素)。不同小写字母表示差异显著(P < 0.05)。

Fig. 4 Effects of removal of seed coat surrounding embryo on seed germination and germination of excised embryos. A, Scarified seeds (removed seed coat surrounding embryo) germinated on moistened filter paper; B, Excised embryos germinated on MS medium (no exogenous phytohormone). Different letters show significant difference at P = 0.05 level.

图5 不同种源的葡萄品种种子在不同温度条件下的层积反应。(A) 东亚种; (B) 欧亚种; (C) 美洲种、欧山杂种、蘡欧杂种、欧美杂种在5°C、25°C、20/10°C和30/20°C下层积2个月后的萌发率; (D) 美洲种和欧美杂种在5°C和20/10°C下层积6个月后的萌发率。

Fig. 5 Stratification effects at different temperatures on seed germination of grape varieties from different proveniences. (A) Varieties of East-Asia species; (B) Europe-Asia species; (C) Varieties of American species, cross-breeds of Europe-Asia species × East-Asia species, East-Asia species × Europe-Asia species, Europe-Asia species × American species; A, B, C: stratified for 2 months at 5°C, 25°C, 20/10°C and 30/20°C. (D) Varieties of American species and Europe-Asia species × American species, stratified for 6 months at 5°C and 20/10°C.

图6 不同种源的葡萄种子在模拟环境四季平均温度变化下的萌发进程。循环1(●): 夏季 (3个月, 30/20°C)—秋季 (3个月, 20/10°C)—冬季 (3个月, 5°C)—春季 (3个月, 20/10°C); 循环2 (○): 冬季 (3个月, 5°C)—春季 (3个月, 20/10°C)—夏季 (3个月, 30/20°C)—秋季(3个月, 20/10°C)。箭头表示种子从一个温度范围转移到下一个温度范围的季节转化。

Fig. 6 Germination course of different species of grape seeds at alternate temperatures simulating the season changes. Cycle 1(●): Summer (3 months, 30/20°C)-autumn (3 months, 20/10°C)-winter (3 months, 5°C)-spring (3 months, 20/10°C); Cycle 2 (○): Winter (3 months, 5°C)-spring (3 months, 20/10°C)-summer (3 months, 30/20°C)-autumn (3 months, 20/10°C). Arrows indicate the simulating season changes at which seeds were moved from one temperature regime to the next.

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