杨树品系低温生理响应及相关差异蛋白分析
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摘要
杨树是我国北方重要的造林及城乡绿化树种,其冻害频繁发生及由冻害引发的病害是杨树产业发展的重要制约因素之一。因此,研究秋季自然低温胁迫下不同杨树品系(种)生理生化响应机制及差异蛋白的表达特性,对了解杨树抗寒调节机理具有重要的理论意义,对如何判别现有杨树品种在其栽培地点对低温变化的适应性；如何在育种中平衡生长量和适应性进行杂交亲本选配,以及杂种无性系对比试验和区域化试验地点的选择等生产实践问题具有一定指导作用。
本研究选择适生区域不同的大青杨、小黑杨、黑林5号杨、1333杨等杨树品系(种)为研究材料,在哈尔滨地区就秋季低温胁迫下不同杨树品系(种)生长、物候、冻害发生,以及叶片细胞膜透性、叶绿素荧光参数、渗透调节物质、酚类物质、特异蛋白等变化规律,以探讨秋季自然低温胁迫下杨树抗寒响应机制。研究结果表明：
(1)不同杨树品系(种)在同一栽培地点对低温变化的反应不同,遗传基础决定其生长、物候以及适应性差异。其中,大青杨为当地乡土树种,在其长期进化过程中适应了当地的气候特点,其封顶最早,顶芽越冬率最高,基本无冻害问题；黑林5号杨封顶略晚于大青杨,顶芽越冬率(97%)次之：小黑杨封顶晚于黑林5号杨,顶芽越冬率也随之下降(90.5%)；1333杨封顶最晚,甚至未形成休眠顶芽,生长周期长,树叶宿存直至冻落,其光周期与当地气候适应性差,难以通过适时休眠规避低温伤害。
(2)平均最低气温5℃时,相对电导率与叶绿素荧光参数表明杨树叶片遭受可逆胁迫,说明这两者能反映杨树受低温胁迫程度的重要生理指标。零上低温胁迫,大青杨表现受胁迫最重,其次为黑林5号杨与小黑杨,1333杨受害最轻,气温降至零下3.3℃,4个杨树品系(种)相对电导率均超过50%,大青杨表现为受害最轻,1333杨受害最严重,这表明零上低温时代谢旺盛的1333杨耐受低温胁迫能力强,零下低温时,及时休眠可能是杨树应对低温的更好方式。
(3)遭受低温胁迫时杨树叶片中渗透调节物质与可溶性酚类物质表现不同程度上升,表明这些保护物质与杨树抗寒性关系密切。1333杨响应最强烈,其叶片中渗透调节物质、可溶性酚类物质随气温下降而上升,多表现为显著性负相关,其次是小黑杨与黑林5号杨,大青杨叶片中渗透调节物质与酚类物质多数对零上低温有一定响应,落叶前都出现大幅下降。这是不同品系(种)在低温胁迫中不同抗寒策略的体现,大青杨主要通过避逆的方式应对低温胁迫,1333杨主要通过提高保护物质含量来响应低温胁迫。
(4)大青杨封顶早,落叶前渗透调节物质大量转移,电导率、Fv/Fm、Fv/F0在零下低温来临前平缓下降等都是大青杨在长期进化过程中与环境相互适应的一种表现,在哈尔滨地区适应性较好的小黑杨、黑林5号杨电导率、Fv/Fm、Fv/F0在零下低温来临前也表现为平缓下降,叶绿素荧光参数的测定具有快速、无伤害等优点,因此可以作为杨树生态适应性的生理指标继续进行研究。
(5)低温胁迫对杨树蛋白质表达有显著影响,在pH 4～7范围内,大青杨蛋白质双向电泳表达图谱中发现94个蛋白质点在秋季低温胁迫下有丰度变化2倍以上,45个蛋白质被成功鉴定,GO分类结果表明23个(42%)蛋白是胁迫响应蛋白,12个(23%)蛋白与呼吸作用相关,9个(17%)蛋白与光合作用相关,5个(10%)蛋白参与其它生理过程,还有4个(8%)蛋白功能未知。鉴定的45个蛋白中,有41个表达量上调,4个表达量下调。这些蛋白表达丰度变化可能与大青杨特有的抗寒适应机制密切相关。
(6)以大青杨凝胶图谱上已鉴定的45个蛋白为参照,将小黑杨、1333杨3个时期分别与大青杨凝胶图谱进行匹配,28个蛋白点与小黑杨、1333杨匹配良好,定量分析后与大青杨中同一蛋白点进行比较,结果15个蛋白点在3个品系(种)中表达量均上调,13个蛋白点变化规律不同,其中9个蛋白点在大青杨叶片中表达量上调,小黑杨、1333杨叶片中上调不显著,另外4个蛋白点在大青杨叶片中表达量下调,小黑杨、1333杨叶片中表达丰度变化未超过2倍。变化规律不同的蛋白可能与3个杨树品系(种)低温适应机制不同密切相关,有待于进一步研究。
(7)热激蛋白(点4、10、67)、磷酸甘油酸变位酶(点21)、细胞质蛋白动力蛋白重链(点44)、ATP合酶δ链(点53)6个蛋白点在3个杨树品系(种)中表达差异可能与其应对低温胁迫机制密切相关。
Poplar is an important afforestation and greening tree species in Northern China. Frequent cold injury and the triggered disease are mainly limited factors for poplar industrial development. Therefore, it is necessary to study physiological and biochemical response of different cultivars to low temperature in autumn and specific expression of function proteins. It has the significance to elaborate the mechanism of cold resistance, distinguish the adaptability of different cultivars, and keep the balance of mass growth and adaptability for the selective breeding of cross parent, as well as select the sites for contrast test and compartmentation test of hybridal clones.
Four poplar cultivars [P. ussuriensis Kom,populus simonii×P. nigra, (P. psudosimonii×P. nigra)×P. nigra CL.'A5'and populus alba×P. davidian CL.'1333'] grown in Harbin city were selected to investigate their cold resistance to natural low temperature in autumn; cold-resistance mechanisms were studied by investigating on the changes of growth, phenology, cell membrane permeability, chlorophyll fluorescence parameters, osmotic adjustment, phenolic compounds and changes of specific proteins in natural cooling process. The results were shown as follows:
(1) Different poplar cultivars grown in the same site had various resistances to the changing temperature; genetic basis was a determining factor for the difference in growth, phenology, and adaptability. P. ussuriensis, belonging to the local native tree species, forms some adaptive characteristics to local climate in a long-term evolution process, such as earlier occurrence of capping and higher survival rate of terminal buds, thus this cultivar had almost no cold injure to occur. (P. psudosimonii×P. nigra)×P. nigra CL.'A5'cap slightly later than that of P. ussuriensis, its survival rate of the terminal buds in winter was 97%; the capping of Populus simonii×P. nigra and populus alba×P. davidian CL.'1333'occurred later; this led to the difficulty of the formation of dormant buds and a poor adaptability of its photoperiod to local climate. Thus the two cultivars could not avoid chilling injury by timely access dormant.
(2) Under the average minimum temperature of 5℃, significant changes in relative conductivity and chlorophyll fluorescence parameters were observed in the four cultivars. Under the stress of low temperature above zero, the injured degree of P. ussuriensis was the highest, followed by (P. psudosimonii×P. nigra)×P. nigra CL.'A5', Populus simonii×P. nigra and populus alba XP. davidian CL.'1333'. However, when air temperature dropped to-3℃, relative conductivity of the four poplar cultivars exceeded 50%, of which the injured degree of P. ussuriensis was the lowest, populus alba×P. davidian CL.'1333'the highest. It indicates that timely access dormant may be a better way to resist low temperature for the poplar cultivars.
(3) Under chilling stress, osmotic regulation substances and phenolic compounds of the poplar leaves were increased, indicating that there is a close correlation between protection substances and cold resistance. In general, osmotic regulation substances and phenolic compounds of in leaves were increased with a decreasing temperature. The strongest response was found in populus alba×P. davidian CL.'1333'leaves, followed by populus simonii×P. nigra and(P. psudosimonii×P. nigra)×P. nigra CL.'A5'. A relatively weak response occurred only at low temperature above zero for P. ussuriensis. A large decrease in osmotic regulation substances and phenolic compounds occurred for all the four cultivars before defoliation. It implies that P. ussuriensis adapts the low temperature stress mainly through the way of stress avoidance, whereas populus alba×P. davidian CL.'1333'resists the chilling stress by improving protective substances.
(4) P. ussuriensis had an earlier capping period. Before defoliation, osmotic regulation substances were transferred largely between organs, moreover, relative conductivity, Fv/Fm, and Fv/F0 were decreased slowly before freezing temperature arrival. These characteristics are adaptive performance, formed in a long-term evolution. The other two cultivars (populus simonii×P. nigra and(P. psudosimonii×P. nigra)×P. nigra CL.'A5') also showed a similar changing trend.
(5) Low temperature stress had significant effects on the protein expression. Under the condition of pH 4-7,94 protein spots in gel electrophoresis map of P. ussuriensis were found to have the change in abundance of more than two times, of which 45 spots were identified. GO classification showed that 23 spots (42% of total protein spots) are stress response proteins,12 (23%) spots might be related to energy metabolism, nine spots (17%) related to photosynthesis, five spots (10%) catabolism associated, the rest four spots (8%) unidentified for the function. Among the identified 45 proteins,41 spots were up-regulated expression; only four spots were down-regulated. These changes in protein abundance may be closely related to chilling adaptation mechanisms in P. ussuriensis.
(6) Compared with the 45 protein spots identified in gel electrophoresis map of P. ussuriensis,28 protein spots were matched in gel electrophoresis map of populus simonii×P. nigra and populus alba X P. davidian CL.'1333'. Among them, the expressions of 15 proteins were up-regulated in three cultivars in the three stages of temperature drop, the other 13 protein spots showed an inregulatory express, of which nine proteins in P. ussuriensis were up-regulated expression and four proteins were down-regulated expression; the change in populus simonii×P. nigra and populus alba×P. davidian CL. '1333'are not significantly. Variation in protein expression in the three cultivars may be closely related with different low-temperature adaptation mechanisms, this assumption needs to be studied further.
(7)The expression differences of heat shock protein (spot 4,10,67), bisphosphoglycerate-independent phosphoglycerate mutase (spot 21), filamentation temperature-sensitive h 2b (spot 44), ATP synthase delta chain (spot 53) in the three poplar species may be closely related to low temperature stress response mechanisms.
本研究选择适生区域不同的大青杨、小黑杨、黑林5号杨、1333杨等杨树品系(种)为研究材料,在哈尔滨地区就秋季低温胁迫下不同杨树品系(种)生长、物候、冻害发生,以及叶片细胞膜透性、叶绿素荧光参数、渗透调节物质、酚类物质、特异蛋白等变化规律,以探讨秋季自然低温胁迫下杨树抗寒响应机制。研究结果表明：
(1)不同杨树品系(种)在同一栽培地点对低温变化的反应不同,遗传基础决定其生长、物候以及适应性差异。其中,大青杨为当地乡土树种,在其长期进化过程中适应了当地的气候特点,其封顶最早,顶芽越冬率最高,基本无冻害问题；黑林5号杨封顶略晚于大青杨,顶芽越冬率(97%)次之：小黑杨封顶晚于黑林5号杨,顶芽越冬率也随之下降(90.5%)；1333杨封顶最晚,甚至未形成休眠顶芽,生长周期长,树叶宿存直至冻落,其光周期与当地气候适应性差,难以通过适时休眠规避低温伤害。
(2)平均最低气温5℃时,相对电导率与叶绿素荧光参数表明杨树叶片遭受可逆胁迫,说明这两者能反映杨树受低温胁迫程度的重要生理指标。零上低温胁迫,大青杨表现受胁迫最重,其次为黑林5号杨与小黑杨,1333杨受害最轻,气温降至零下3.3℃,4个杨树品系(种)相对电导率均超过50%,大青杨表现为受害最轻,1333杨受害最严重,这表明零上低温时代谢旺盛的1333杨耐受低温胁迫能力强,零下低温时,及时休眠可能是杨树应对低温的更好方式。
(3)遭受低温胁迫时杨树叶片中渗透调节物质与可溶性酚类物质表现不同程度上升,表明这些保护物质与杨树抗寒性关系密切。1333杨响应最强烈,其叶片中渗透调节物质、可溶性酚类物质随气温下降而上升,多表现为显著性负相关,其次是小黑杨与黑林5号杨,大青杨叶片中渗透调节物质与酚类物质多数对零上低温有一定响应,落叶前都出现大幅下降。这是不同品系(种)在低温胁迫中不同抗寒策略的体现,大青杨主要通过避逆的方式应对低温胁迫,1333杨主要通过提高保护物质含量来响应低温胁迫。
(4)大青杨封顶早,落叶前渗透调节物质大量转移,电导率、Fv/Fm、Fv/F0在零下低温来临前平缓下降等都是大青杨在长期进化过程中与环境相互适应的一种表现,在哈尔滨地区适应性较好的小黑杨、黑林5号杨电导率、Fv/Fm、Fv/F0在零下低温来临前也表现为平缓下降,叶绿素荧光参数的测定具有快速、无伤害等优点,因此可以作为杨树生态适应性的生理指标继续进行研究。
(5)低温胁迫对杨树蛋白质表达有显著影响,在pH 4～7范围内,大青杨蛋白质双向电泳表达图谱中发现94个蛋白质点在秋季低温胁迫下有丰度变化2倍以上,45个蛋白质被成功鉴定,GO分类结果表明23个(42%)蛋白是胁迫响应蛋白,12个(23%)蛋白与呼吸作用相关,9个(17%)蛋白与光合作用相关,5个(10%)蛋白参与其它生理过程,还有4个(8%)蛋白功能未知。鉴定的45个蛋白中,有41个表达量上调,4个表达量下调。这些蛋白表达丰度变化可能与大青杨特有的抗寒适应机制密切相关。
(6)以大青杨凝胶图谱上已鉴定的45个蛋白为参照,将小黑杨、1333杨3个时期分别与大青杨凝胶图谱进行匹配,28个蛋白点与小黑杨、1333杨匹配良好,定量分析后与大青杨中同一蛋白点进行比较,结果15个蛋白点在3个品系(种)中表达量均上调,13个蛋白点变化规律不同,其中9个蛋白点在大青杨叶片中表达量上调,小黑杨、1333杨叶片中上调不显著,另外4个蛋白点在大青杨叶片中表达量下调,小黑杨、1333杨叶片中表达丰度变化未超过2倍。变化规律不同的蛋白可能与3个杨树品系(种)低温适应机制不同密切相关,有待于进一步研究。
(7)热激蛋白(点4、10、67)、磷酸甘油酸变位酶(点21)、细胞质蛋白动力蛋白重链(点44)、ATP合酶δ链(点53)6个蛋白点在3个杨树品系(种)中表达差异可能与其应对低温胁迫机制密切相关。
Poplar is an important afforestation and greening tree species in Northern China. Frequent cold injury and the triggered disease are mainly limited factors for poplar industrial development. Therefore, it is necessary to study physiological and biochemical response of different cultivars to low temperature in autumn and specific expression of function proteins. It has the significance to elaborate the mechanism of cold resistance, distinguish the adaptability of different cultivars, and keep the balance of mass growth and adaptability for the selective breeding of cross parent, as well as select the sites for contrast test and compartmentation test of hybridal clones.
Four poplar cultivars [P. ussuriensis Kom,populus simonii×P. nigra, (P. psudosimonii×P. nigra)×P. nigra CL.'A5'and populus alba×P. davidian CL.'1333'] grown in Harbin city were selected to investigate their cold resistance to natural low temperature in autumn; cold-resistance mechanisms were studied by investigating on the changes of growth, phenology, cell membrane permeability, chlorophyll fluorescence parameters, osmotic adjustment, phenolic compounds and changes of specific proteins in natural cooling process. The results were shown as follows:
(1) Different poplar cultivars grown in the same site had various resistances to the changing temperature; genetic basis was a determining factor for the difference in growth, phenology, and adaptability. P. ussuriensis, belonging to the local native tree species, forms some adaptive characteristics to local climate in a long-term evolution process, such as earlier occurrence of capping and higher survival rate of terminal buds, thus this cultivar had almost no cold injure to occur. (P. psudosimonii×P. nigra)×P. nigra CL.'A5'cap slightly later than that of P. ussuriensis, its survival rate of the terminal buds in winter was 97%; the capping of Populus simonii×P. nigra and populus alba×P. davidian CL.'1333'occurred later; this led to the difficulty of the formation of dormant buds and a poor adaptability of its photoperiod to local climate. Thus the two cultivars could not avoid chilling injury by timely access dormant.
(2) Under the average minimum temperature of 5℃, significant changes in relative conductivity and chlorophyll fluorescence parameters were observed in the four cultivars. Under the stress of low temperature above zero, the injured degree of P. ussuriensis was the highest, followed by (P. psudosimonii×P. nigra)×P. nigra CL.'A5', Populus simonii×P. nigra and populus alba XP. davidian CL.'1333'. However, when air temperature dropped to-3℃, relative conductivity of the four poplar cultivars exceeded 50%, of which the injured degree of P. ussuriensis was the lowest, populus alba×P. davidian CL.'1333'the highest. It indicates that timely access dormant may be a better way to resist low temperature for the poplar cultivars.
(3) Under chilling stress, osmotic regulation substances and phenolic compounds of the poplar leaves were increased, indicating that there is a close correlation between protection substances and cold resistance. In general, osmotic regulation substances and phenolic compounds of in leaves were increased with a decreasing temperature. The strongest response was found in populus alba×P. davidian CL.'1333'leaves, followed by populus simonii×P. nigra and(P. psudosimonii×P. nigra)×P. nigra CL.'A5'. A relatively weak response occurred only at low temperature above zero for P. ussuriensis. A large decrease in osmotic regulation substances and phenolic compounds occurred for all the four cultivars before defoliation. It implies that P. ussuriensis adapts the low temperature stress mainly through the way of stress avoidance, whereas populus alba×P. davidian CL.'1333'resists the chilling stress by improving protective substances.
(4) P. ussuriensis had an earlier capping period. Before defoliation, osmotic regulation substances were transferred largely between organs, moreover, relative conductivity, Fv/Fm, and Fv/F0 were decreased slowly before freezing temperature arrival. These characteristics are adaptive performance, formed in a long-term evolution. The other two cultivars (populus simonii×P. nigra and(P. psudosimonii×P. nigra)×P. nigra CL.'A5') also showed a similar changing trend.
(5) Low temperature stress had significant effects on the protein expression. Under the condition of pH 4-7,94 protein spots in gel electrophoresis map of P. ussuriensis were found to have the change in abundance of more than two times, of which 45 spots were identified. GO classification showed that 23 spots (42% of total protein spots) are stress response proteins,12 (23%) spots might be related to energy metabolism, nine spots (17%) related to photosynthesis, five spots (10%) catabolism associated, the rest four spots (8%) unidentified for the function. Among the identified 45 proteins,41 spots were up-regulated expression; only four spots were down-regulated. These changes in protein abundance may be closely related to chilling adaptation mechanisms in P. ussuriensis.
(6) Compared with the 45 protein spots identified in gel electrophoresis map of P. ussuriensis,28 protein spots were matched in gel electrophoresis map of populus simonii×P. nigra and populus alba X P. davidian CL.'1333'. Among them, the expressions of 15 proteins were up-regulated in three cultivars in the three stages of temperature drop, the other 13 protein spots showed an inregulatory express, of which nine proteins in P. ussuriensis were up-regulated expression and four proteins were down-regulated expression; the change in populus simonii×P. nigra and populus alba×P. davidian CL. '1333'are not significantly. Variation in protein expression in the three cultivars may be closely related with different low-temperature adaptation mechanisms, this assumption needs to be studied further.
(7)The expression differences of heat shock protein (spot 4,10,67), bisphosphoglycerate-independent phosphoglycerate mutase (spot 21), filamentation temperature-sensitive h 2b (spot 44), ATP synthase delta chain (spot 53) in the three poplar species may be closely related to low temperature stress response mechanisms.
引文
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