盐胁迫对欧洲赤松光合作用的影响及耐盐性评价
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摘要
本研究所用的欧洲赤松(Pinus silvestris L.)适应性非常强,常常生长在盐碱土上(Cl- 0.213%, SO42- 0.606%, Na+0.419%),是非常珍贵的耐盐树种。为了进一步丰富我国欧洲赤松种质资源,为盐碱地区选择优良树种,为杂交选育提供优良种源,有必要开展欧洲赤松的引种研究。
本研究以引进的7个欧洲赤松种源和两个国内樟子松种源2年生幼苗为材料,采用大棚盆栽的方法,选取生长指标(苗高、基径)、光合指标,进行耐盐性分析。9个种源分2批种植在不同盐度处理下：(1)哈一、哈二、754、17330、N8、11、17274、大兴安岭、红花尔基,盐处理为0.5%和CK。于胁迫后第10、30、45、60天测量2个生长指标(苗高和基径),第2、20、50天测净光合速率(Pn),第5、30、50天测Fv/Fm；(2)哈一和哈二盐处理梯度为0.25%,0.5%,1%和CK。胁迫后第2、20、50天测净光合速率(Pn)等光合生理指标,胁迫后第25天测光合日变化,第2、20、50天时测光响应曲线,第5、30、60天测PSⅡ叶绿素荧光参数。
主要结果为：
1.选取了苗高、基径、相净光合速率、Fv/Fm相对变化率这4个指标,对引进的7个欧洲赤松种源和两个国内樟子松种源进行耐盐性评价,评价结果为：哈一>哈二>17330、11>754>N8>17274>红花尔基>大兴安岭。引进的7个种源的耐盐性都高于国内红花尔基和大兴安岭两个种源的耐盐性,哈一和哈二两个种源的耐盐性排在前两位。
2.根据Pn、Gs、Ci、Tr、Vpdl、Ls、WUE、CE测量结果表明：盐胁迫导致哈一、哈二的Pn的降低既有浓度效应又有时间效应；盐胁迫导致Pn降低的因素,哈一为气孔限制因素；哈二对短期的盐胁迫表现为气孔限制因素,而对长期的盐胁迫表现为气孔限制因素和非气孔限制因素共同作用。通过回归分析,结果显示盐处理后,影响哈一Pn的光合生态因子排列顺序为CE>Ci>Gs>WUE>Ls>Vpdl,影响哈二Pn的光合生态因子的排列顺序为,Ci>CE>Ls>Gs>Tr>WUE。
3.根据Pn、GS、Ci、Tr、Vpdl、Ls、WUE、LUE、CE的日变化结果表明：哈一在0.25%盐处理时Pn日变化为单峰曲线,CK、0.5%盐处理时Pn日变化为为双峰曲线,1%盐处理时Pn日变化为波浪形曲线；影响哈-CK Pn日变化的生态因子排序为RH>Ci>Ca >Ta>Gs>Vpdl; 0.25%盐处理影响Pn日变化的生态因子排序为RH>Tr>Ci>Ca>Ta>PAR：0.5%盐处理影响Pn日变化的生态因子排序为Ta>RH>Vpdl>Ci>PAR;1%盐处理影响Pn日变化的生态因子排序为Ci>Ca>Gs。哈二CK和0.25%盐处理时Pn日变化为单峰曲线,0.5%和1%盐处理时Pn日变化为双峰曲线；影响哈二CK Pn日变化的生态因子排序为Ci>Vpdl>RH>Tr>Ca>Gs; 0.25%盐处理影响P。日变化的生态因子排序为Vpdl>Ta> RH>Ci>PAR>Tr; 0.5%盐处理影响Pn日变化的生态因子排序为Ca>Ci>PAR>Tr>Ta；1%盐处理影响Pn日变化的生态因子排序为Tr>Ci>PAR>Ca>Ta>RHo
4.光响应曲线拟合结果表明：模型5对欧洲赤松哈一和哈二ck、0.25%、0.5%、1%的拟合结果要好于其他模型。哈一和哈二在盐处理后AQY和LSP降低,可能是造成P。降低的原因。
5.Fv/Fm、Fv'/Fm'、ΦPSⅡ、PhiCO2、qP、qN、ETR测量结果表明：哈一盐处理后Fv/Fm、Fv'/Fm'、ΦPSⅡ、ETR、PhiCO2与盐胁迫强度呈极显著负相关,qN与盐胁迫强度呈极显著正向相关,qP与盐胁迫强度呈不显著负相关,盐胁下PSⅡ光合反应中心受到损害、表观电子传递速率降低以及光合电子传递受阻是导致哈一Pn降低的一个重要原因。哈二在盐处理后Fv/Fm、Fv'/Fm'、ΦPSⅡ、qP、ETR、PhiCO2与盐胁迫都出极显著负相关,qN与盐胁迫强度呈极显著正相关,盐胁迫下PSⅡ光合反应中心受到损害以及CO2同化速率相对应的量子产量降低是导致哈二Pn降低的一个重要原因。
盐胁迫下,哈一和哈二的光合作用都受到不同程度抑制,哈二P。降低幅度,Pmax降低的幅度,WUE、LUE、CE日平均值降低幅度,日碳同化量降低的幅度、PSⅡ光合反应中心受到损害程度都大于哈一,说明哈一的耐盐性优于哈二。
Pinus silvestris L., an adaptability precious salt-tolerance variety, could grow on the saline-alkali soil (Cl- 0.213%, SO42- 0.606%, Na+ 0.419%). A study on introduction of Pinus silvestris L. is necessary to be carried out in order to enriching genetic resources of Pinus silvestris L., selecting excellent species, and providing crossing parents in China.
In a pot experiment made by saline-alkali soil, this paper studied the salt-tolerance of 7 introduced Pinus silvestris L. provenances and 2 Pinus sylvestris var. mongolica domestic ones. The 9 kind of 2-year-old seedlings were divide into 2 batches to plant:(1) Hal, Ha2,754, 17330, N8,11,17274, DAXINGANLING, HONGHUAERJI, the salt stress level is 0.5% and control. After salt treatment,2 grow indicators, height growth and stem basal diameter, were determined on the 10th,30th,45th,60th day. Net photosynthetic rate (Pn) measured on the 2nd, 20th,50th day and Fv/Fm measured on the 5th,30th,50th day. (2) Hal, Ha2, the salt stress level is 0.25%,0.5%,1.0% and control. After salt treatment, diurnal variation of 9 photosynthesis indexes was measured on the 25th day. Photosynthesis physiological indexes and Light responsive curve measured on the 2nd,20th,50th day and chlorophyll fluorescence parameters measured on the 5th,30th,60th day. The mainly conclusions as follow:
1. Four indexes, relative variable ratio of height growth, basal diameter, Pn and Fv/Fm were used to evaluate the salt tolerance of 7 introduced Pinus silvestris L. provenances and 2 Pinus sylvestris var. mongolica domestic ones, its order is:Hal>Ha2>17330 and 11>754>N8> 17274>HONGHUAERJI>DAXINGANLING.7 introduced Pinus silvestris L. provenances have stronger tolerance to salt stress than those of 2 domestic ones (HONGHUAERJI and DAXINGANLING), tolerance to salt stress of Hal and Ha2 was superior.
2. According to the results of Pn, Gs, Ci, Tr, Vpdl, Ls, WUE and CE, descended Pn under salt stress of Hal and Ha2 has both concentration effects and time effects. It denoted the main factor inhibiting Pn of Hal is stomatal control. The factor inhibiting Pn of Ha2 to short-time salt stress is stomatal control, and that to long-time salt stress is stomata and non-stomatal control. Regression analysis results showed that effects of salt stress on photosynthetic ecological factors of Hal order is CE>Ci>Gs> WUE>Ls> Vpdl, and that of Ha2 is Ci>CE >LS>GS>Tr>WUE.
3. According to the results of diurnal variation of Pn, Gs, Ci, Tr, Vpdl, Ls, WUE, LUE and CE, the Pn diurnal variation curves of Hal under 0.25%, CK together with 0.5%,1% salt stress is single-peak, two-peak and waves respectively. Ecological factors orders of Hal under CK, 0.25%,0.5% and 1% salt treatments respectively is RH>Ci>Ca>Ta>Gs>Vpdl, RH>Tr>Ci >Ca>Ta>PAR, Ta>RH>Vpdl>Ci>PAR, Ci>Ca>GS. The Pn diurnal variation curves of Hal under CK together with 0.25%,0.5% together with 1% salt stress is single-peak and two-peak respectively. Ecological factors orders of Hal under CK,0.25%,0.5% and 1% salt treatments respectively is Ci>Vpdl>RH>Tr>Ca>Gs, Vpdl>Ta>RH>Ci>PAR>T, Ca>Ci >PAR>Tr>Ta, Tr>Ci>PAR>Ca>Ta>RH.
4. The light response curves fitting results showed that model 5 were better than the others. Descended AQYand LSP might be important factors resulting in the decrease of Pn.
5. According to the results of Fv/Fm, Fv'/Fm',ΦPSⅡ, PhiCO2, qP, qN, ETR, under salt stress, the statistics analysis of Hal showed a significant negative correlation between salinity and Fv/Fm, Fv'/Fm',ΦPSⅡ, ETR, PhiCO2, a significant positive correlation between salinity and qN, but a non-significant negative correlation between salinity and qP. The reaction center inactivation, reducing electrons transport rate and hampering photosynthetic electron transfer might be important factors resulting in the decrease Pn of Hal. The statistics analysis of Ha2 showed a significant negative correlation between salinity and Fv/Fm, Fv'/Fm',ΦPSⅡ, ETR, qp, PhiCO2, a significant positive correlation between salinity and qN. The reaction center inactivation, reducing quantum yield corresponding to CO2 assimilation rate might be important factors resulting in the decrease Pn of Ha2.
Under salt stress, photosynthesis was inhibited in varying degrees. The Ha2 value of decreased Pn and Pmax, daily average of WUE, LUE, CE, damage of the reaction center were greater than those of Hal. Overall, the salt tolerance of Hal is superior to Ha2.
本研究以引进的7个欧洲赤松种源和两个国内樟子松种源2年生幼苗为材料,采用大棚盆栽的方法,选取生长指标(苗高、基径)、光合指标,进行耐盐性分析。9个种源分2批种植在不同盐度处理下：(1)哈一、哈二、754、17330、N8、11、17274、大兴安岭、红花尔基,盐处理为0.5%和CK。于胁迫后第10、30、45、60天测量2个生长指标(苗高和基径),第2、20、50天测净光合速率(Pn),第5、30、50天测Fv/Fm；(2)哈一和哈二盐处理梯度为0.25%,0.5%,1%和CK。胁迫后第2、20、50天测净光合速率(Pn)等光合生理指标,胁迫后第25天测光合日变化,第2、20、50天时测光响应曲线,第5、30、60天测PSⅡ叶绿素荧光参数。
主要结果为：
1.选取了苗高、基径、相净光合速率、Fv/Fm相对变化率这4个指标,对引进的7个欧洲赤松种源和两个国内樟子松种源进行耐盐性评价,评价结果为：哈一>哈二>17330、11>754>N8>17274>红花尔基>大兴安岭。引进的7个种源的耐盐性都高于国内红花尔基和大兴安岭两个种源的耐盐性,哈一和哈二两个种源的耐盐性排在前两位。
2.根据Pn、Gs、Ci、Tr、Vpdl、Ls、WUE、CE测量结果表明：盐胁迫导致哈一、哈二的Pn的降低既有浓度效应又有时间效应；盐胁迫导致Pn降低的因素,哈一为气孔限制因素；哈二对短期的盐胁迫表现为气孔限制因素,而对长期的盐胁迫表现为气孔限制因素和非气孔限制因素共同作用。通过回归分析,结果显示盐处理后,影响哈一Pn的光合生态因子排列顺序为CE>Ci>Gs>WUE>Ls>Vpdl,影响哈二Pn的光合生态因子的排列顺序为,Ci>CE>Ls>Gs>Tr>WUE。
3.根据Pn、GS、Ci、Tr、Vpdl、Ls、WUE、LUE、CE的日变化结果表明：哈一在0.25%盐处理时Pn日变化为单峰曲线,CK、0.5%盐处理时Pn日变化为为双峰曲线,1%盐处理时Pn日变化为波浪形曲线；影响哈-CK Pn日变化的生态因子排序为RH>Ci>Ca >Ta>Gs>Vpdl; 0.25%盐处理影响Pn日变化的生态因子排序为RH>Tr>Ci>Ca>Ta>PAR：0.5%盐处理影响Pn日变化的生态因子排序为Ta>RH>Vpdl>Ci>PAR;1%盐处理影响Pn日变化的生态因子排序为Ci>Ca>Gs。哈二CK和0.25%盐处理时Pn日变化为单峰曲线,0.5%和1%盐处理时Pn日变化为双峰曲线；影响哈二CK Pn日变化的生态因子排序为Ci>Vpdl>RH>Tr>Ca>Gs; 0.25%盐处理影响P。日变化的生态因子排序为Vpdl>Ta> RH>Ci>PAR>Tr; 0.5%盐处理影响Pn日变化的生态因子排序为Ca>Ci>PAR>Tr>Ta；1%盐处理影响Pn日变化的生态因子排序为Tr>Ci>PAR>Ca>Ta>RHo
4.光响应曲线拟合结果表明：模型5对欧洲赤松哈一和哈二ck、0.25%、0.5%、1%的拟合结果要好于其他模型。哈一和哈二在盐处理后AQY和LSP降低,可能是造成P。降低的原因。
5.Fv/Fm、Fv'/Fm'、ΦPSⅡ、PhiCO2、qP、qN、ETR测量结果表明：哈一盐处理后Fv/Fm、Fv'/Fm'、ΦPSⅡ、ETR、PhiCO2与盐胁迫强度呈极显著负相关,qN与盐胁迫强度呈极显著正向相关,qP与盐胁迫强度呈不显著负相关,盐胁下PSⅡ光合反应中心受到损害、表观电子传递速率降低以及光合电子传递受阻是导致哈一Pn降低的一个重要原因。哈二在盐处理后Fv/Fm、Fv'/Fm'、ΦPSⅡ、qP、ETR、PhiCO2与盐胁迫都出极显著负相关,qN与盐胁迫强度呈极显著正相关,盐胁迫下PSⅡ光合反应中心受到损害以及CO2同化速率相对应的量子产量降低是导致哈二Pn降低的一个重要原因。
盐胁迫下,哈一和哈二的光合作用都受到不同程度抑制,哈二P。降低幅度,Pmax降低的幅度,WUE、LUE、CE日平均值降低幅度,日碳同化量降低的幅度、PSⅡ光合反应中心受到损害程度都大于哈一,说明哈一的耐盐性优于哈二。
Pinus silvestris L., an adaptability precious salt-tolerance variety, could grow on the saline-alkali soil (Cl- 0.213%, SO42- 0.606%, Na+ 0.419%). A study on introduction of Pinus silvestris L. is necessary to be carried out in order to enriching genetic resources of Pinus silvestris L., selecting excellent species, and providing crossing parents in China.
In a pot experiment made by saline-alkali soil, this paper studied the salt-tolerance of 7 introduced Pinus silvestris L. provenances and 2 Pinus sylvestris var. mongolica domestic ones. The 9 kind of 2-year-old seedlings were divide into 2 batches to plant:(1) Hal, Ha2,754, 17330, N8,11,17274, DAXINGANLING, HONGHUAERJI, the salt stress level is 0.5% and control. After salt treatment,2 grow indicators, height growth and stem basal diameter, were determined on the 10th,30th,45th,60th day. Net photosynthetic rate (Pn) measured on the 2nd, 20th,50th day and Fv/Fm measured on the 5th,30th,50th day. (2) Hal, Ha2, the salt stress level is 0.25%,0.5%,1.0% and control. After salt treatment, diurnal variation of 9 photosynthesis indexes was measured on the 25th day. Photosynthesis physiological indexes and Light responsive curve measured on the 2nd,20th,50th day and chlorophyll fluorescence parameters measured on the 5th,30th,60th day. The mainly conclusions as follow:
1. Four indexes, relative variable ratio of height growth, basal diameter, Pn and Fv/Fm were used to evaluate the salt tolerance of 7 introduced Pinus silvestris L. provenances and 2 Pinus sylvestris var. mongolica domestic ones, its order is:Hal>Ha2>17330 and 11>754>N8> 17274>HONGHUAERJI>DAXINGANLING.7 introduced Pinus silvestris L. provenances have stronger tolerance to salt stress than those of 2 domestic ones (HONGHUAERJI and DAXINGANLING), tolerance to salt stress of Hal and Ha2 was superior.
2. According to the results of Pn, Gs, Ci, Tr, Vpdl, Ls, WUE and CE, descended Pn under salt stress of Hal and Ha2 has both concentration effects and time effects. It denoted the main factor inhibiting Pn of Hal is stomatal control. The factor inhibiting Pn of Ha2 to short-time salt stress is stomatal control, and that to long-time salt stress is stomata and non-stomatal control. Regression analysis results showed that effects of salt stress on photosynthetic ecological factors of Hal order is CE>Ci>Gs> WUE>Ls> Vpdl, and that of Ha2 is Ci>CE >LS>GS>Tr>WUE.
3. According to the results of diurnal variation of Pn, Gs, Ci, Tr, Vpdl, Ls, WUE, LUE and CE, the Pn diurnal variation curves of Hal under 0.25%, CK together with 0.5%,1% salt stress is single-peak, two-peak and waves respectively. Ecological factors orders of Hal under CK, 0.25%,0.5% and 1% salt treatments respectively is RH>Ci>Ca>Ta>Gs>Vpdl, RH>Tr>Ci >Ca>Ta>PAR, Ta>RH>Vpdl>Ci>PAR, Ci>Ca>GS. The Pn diurnal variation curves of Hal under CK together with 0.25%,0.5% together with 1% salt stress is single-peak and two-peak respectively. Ecological factors orders of Hal under CK,0.25%,0.5% and 1% salt treatments respectively is Ci>Vpdl>RH>Tr>Ca>Gs, Vpdl>Ta>RH>Ci>PAR>T, Ca>Ci >PAR>Tr>Ta, Tr>Ci>PAR>Ca>Ta>RH.
4. The light response curves fitting results showed that model 5 were better than the others. Descended AQYand LSP might be important factors resulting in the decrease of Pn.
5. According to the results of Fv/Fm, Fv'/Fm',ΦPSⅡ, PhiCO2, qP, qN, ETR, under salt stress, the statistics analysis of Hal showed a significant negative correlation between salinity and Fv/Fm, Fv'/Fm',ΦPSⅡ, ETR, PhiCO2, a significant positive correlation between salinity and qN, but a non-significant negative correlation between salinity and qP. The reaction center inactivation, reducing electrons transport rate and hampering photosynthetic electron transfer might be important factors resulting in the decrease Pn of Hal. The statistics analysis of Ha2 showed a significant negative correlation between salinity and Fv/Fm, Fv'/Fm',ΦPSⅡ, ETR, qp, PhiCO2, a significant positive correlation between salinity and qN. The reaction center inactivation, reducing quantum yield corresponding to CO2 assimilation rate might be important factors resulting in the decrease Pn of Ha2.
Under salt stress, photosynthesis was inhibited in varying degrees. The Ha2 value of decreased Pn and Pmax, daily average of WUE, LUE, CE, damage of the reaction center were greater than those of Hal. Overall, the salt tolerance of Hal is superior to Ha2.
引文
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