吉林省不同年代大豆品种光合作用与根系活力变化的研究
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摘要
关于大豆品种遗传改良过程中农艺性状和生理变化的研究已有许多报道,但关于根系生理功能变化的研究尚未见报道,根系活力与地上生物量及光合作用的关系目前仍不清楚。本研究通过对吉林省1923年以来育成的主推大豆品种的叶片光合特性和根系伤流量及地上生物量的演化特征,并通过分析光合特性与产量的关系,根系伤流量和根系活力与地上生物量的关系,试图阐明吉林省大豆遗传改良过程中叶片光合特性的演化规律,并揭示根系与地上部分间的相互作用关系,为吉林省大豆品种改良提供可参考的理论依据。
该试验于2010年和2011年两年进行,选择1923年至2009年间育成的大豆品种进行研究,通过对不同年代育成大豆品种叶片光合速率、气孔导度、蒸腾速率、叶绿素和比叶重等光合指标的测定,以及对不同生育期根系伤流量和根系活力及地上和地下生物量的测定,进一步分析其之间的关系,研究结果表明：
(1)从1923年到2005年,82年来每公顷籽粒产量提高80.2%,平均每年增加0.98%。生育期82年来缩短了14.7天,平均每10年缩短1.83天,而产量随大豆品种熟期的缩短而呈增加的变化趋势；收获指数提高了58.03%,且与产量呈极显著正相关(r=0.8099**)；叶片的净光合速率增加了3.9196μmolCO2·m-1·s-1,提高了18.09%,平均每年提高0.22%,产量与叶片净光合速率呈极显著正相关(r=0.6102**)；叶片气孔导度提高了22.91%,平均每年提高0.27%,但产量与叶片气孔导度未达到显著正相关水平；叶片胞间二氧化碳浓度降低了8.3804μmolCO2·mol-1,平均每年降低0.1022μmolCO2·mol-1;蒸腾速率提高了57.29%,平均每年提高0.69%,产量随蒸腾速率的增加而增加,相关系数为0.6316**,而叶片水分利用效率随品种育成年代增加而显著减少,82年来减少了22.3%；叶片表观叶肉导度提高了49.54%,平均每年提高0.60%,产量随表观叶肉导度的增加而显著增加；叶片气孔限制值降低了8.25%,平均每年降低了0.10%,产量随叶片气孔限制值的增加而降低；叶片比叶重提高了91.82%,平均每年提高1.12%,产量随比叶重的增加而增加,相关系数为0.4464*；叶片叶绿素含量提高了17.41%,平均每年提高0.21%,产量随叶绿素的增加而增加；小叶面积降低了47.37%,平均每年降低0.58%。籽粒产量与Pn和Pn/Ci正相关达到极显著水平而叶片的水分利用效率随品种育成年代而显著降低与籽粒产量呈极显著负相关,这三个生理指标可以作为衡量籽粒产量的育种指标。
(2)从根系伤流液和根系活力看,在V4期,根系伤流量86年来增长了9.10%,年平均增长率为0.11%；根系活力增长了10.24%,年平均增长率为0.12%；在R2期,根系伤流量86年来增长了18.29%,年平均增长率为0.21%；根系活力增长了27.38%,年平均增长率为0.32%；在R4期,根系伤流量86年来增长了16.01%,年平均增长率为0.19%；根系活力增长了24.22%,年平均增长率为0.28%；在R6期,伤流液重量86年来增长了20.22%,年平均增长率为0.24%；根系活力增长了10.83%,年平均增长率为0.13%。由此可知,在R4期年增长率达到最大值,同时在R4期伤流液重量与育成年代呈显著正相关,R4期是大豆栽培中保证根系活力的关键阶段,通过优良的栽培措施保证R4期及以后较高的根系活力是保证大豆高产的核心栽培技术手段。
(3)从地上生物量和根系生物量看,在V4期,地上生物量86年来增长了10.81%,年平均增长率为0.13%；根系生物量增长了28.10%,年平均增长率为0.33%；单株生物量增长了11.68%,年平均增长率为0.14%；在R2期,地上部生物量86年来增长了6.99%,年平均增长率为0.08%；根生物量增长了17.65%,年平均增长率为0.21%；单株生物量增长了8.55%,年平均增长率为0.10%；在R4期,地上86年来增长了24.76%,年平均增长率为0.29%；根生物量增长了21.13%,年平均增长率为0.25%；单株生物量增长了24.21%,年平均增长率为0.28%；在R6期,地上部生物量86年来增长了10.51%,年平均增长率为0.12%；根生物量增长了9.27%,年平均增长率为0.11%；单株生物量增长了10.05%,年平均增长率为0.12%。由此可知,根生物量,地上部生物量和单株生物量在R4期与育成年代均呈显著正相关或极显著正相关,在V4、R2和R6期,大豆品种的器官生物量与育成年代虽也呈正相关,但均未达到显著水平。这表明,R4期大豆地上和地下部分的生长对产量起至关重要的作用,大豆品种的遗传改良使植株器官生物量得到显著增加,可能与根系生物量和根系活力的改善有关,特别是R4期的根系活力,在R4期,伤流液重量和根系活力与叶片净光合速率均呈显著正相关,也证明了地下部分与地上光合作用的相关性。因此我们推测根系伤流液重量和根系活力可以作为衡量叶片净光合速率的指标,为今后选育高光合大豆品种提供理论依据。
(4)大豆品种干物质的积累与根系活力、根系伤流液重量有着密切关系。同时我们也发现地上部分与地下根系在品种改良过程中,两者是协同进化的,大豆品种遗传改良是地上器官与地下器官协同进化的结果。在R4期,根系活力和根系伤流液重量和功能叶片的净光合速率关系最为显著,在R2期和R6期,根系活力和叶片净光合速率并未达到显著水平。R2期和R4期地上部分和地下部分有较好的协同关系,但在R6期,根系活力急剧下降,根系伤流液重量迅速减少,但地上部分器官生物量和叶片净光合速率下降不大,说明根系活力的下降先于光合作用的下降,因此大豆植株后期的早衰现象,可能是由于根系的提前衰老导致植株后期光合作用下降的原因。因此我们通过延缓根系的衰老,有可能延缓生育后期光合作用的下降,进而提高产量。
Many reports had showed about agronomic traits and physiological changes in the genetic improvement process of soybean cultivars, but about changes of roots in physiological functions have not been reported, especially about the relationship of root activity and aerial parts of organisms and photosynthesis. This study used soybean varieties bred in Jilin Province since1923to study the photosynthetic characteristics of soybean varieties and root BSW and the biomass of the aboveground, and analyze the relationship of photosynthetic characteristics and yield and the relationship between root BSW and root activity and biomass of aboveground. Aimed to clarify the evolution of the photosynthetic characteristics of leaves in the genetic improvement process of soybean cultivars, and reveal the relationship between the roots and aerial parts, to provide the theoretical basis for the the soybean varieties improvement in Jilin Province.
The experiment was conducted in2010and2011, and chose the soybean varieties bred in the1923-2009. Measured the photosynthetic rate, stomatal conductance, transpiration rate, chlorophyll and specific leaf weight of soybean cultivars, and measured the relationship of root BSW and root activity and biomass of aboveground to analyze their relationship further. The study results show that:
(1) From1923to2005, the grain yield per hectare in82years increased by80.2%, with an average increase by0.98%per year. The of growth period in82years was shortened by14.7days, with an average increase by1.83days every10year, while the yield were showed an increasing trend with shortening in growth period of soybean varieties; harvest index was raise by58.03%, and the harvest index was extremely significant correlated with yield (r=0.8099**); leaf net photosynthetic rate was increased by3.9196μmolCO2·m-1·s-1, and it was increased by18.09%, an average increase was by0.22%per year; the yield was highly significant positive correlated with net photosynthetic rate (r=0.6102**); leaf stomatal conductance was increased by22.91%, an average increase was by0.27%per year, but the correlation of yield and stomatal conductance did not reach significant level; leaf intercellular CO2concentration was decreased by8.3804μmolCO2·mol-1, an average decrease was by0.1022μmolCO2·mol-1per year; transpiration rate was increased by57.29%, an average increase was by0.69%per year, the yield was increased with the increase of transpiration rate, the correlation coefficient was0.6316, but the moisture use efficiency of leaves was decreased significantly along with the year of release, the decrease was by22.3%with the year of release in82years; the blades apparent mesophyll conductance was increased by49.54%, an average increase was by0.60%, the yield was significant increased with the increase in apparent mesophyll conductance; stomatal limitation in leaf was decreased by8.25%, and the average reduction was by0.10%per year, and the yield was decreased with the increase in stomatal limitation; the specific leaf weight was increased by91.82%, an average increase was by1.12%per year, the yield was increased with the increase in specific leaf weight, a correlation coefficient was0.4464; the chlorophyll content in leaf was increased by17.41%, with an average increase was by0.21%per year, the yield was increased with the increase in chlorophyll content; the area of leaflet was decreased by47.37%, with an average reduction was by0.58%. The yield was highly significant positive correlated with Pn, Pn/Ci, but the correlation between the water use efficiency in leaves and year of release was negative, and the water use efficiency in leaves was highly significant negative correlated with grain yield. In concluding, the three physiological indicators could be as indicators by measure of breeding of grain yield.
(2)The analysis on root bleeding sap and root vigor has showed that at V4stage, the root bleeding rate for86years increased by9.10%, the average annual growth rate was0.11%; the root activity grew by10.24%, the average annual growth rate was0.12%; at R2stage, the root bleeding rate increased by18.29%in the86years, the average annual growth rate was0.21%; the root activity grew by27.38%, the average annual growth rate was0.32%; at R4, root injury traffic in1986increased by16.01%, the average annual growth rate was0.19%; root activity grew by24.22%, the average annual growth rate was0.28%; R6of xylem sap weight increase by20.22%in the86years, the average annual growth rate was0.24%; growth of root activity was10.83%, and the average annual growth rate was0.13%. The result showed that at R4, the annual growth rate reached the maximum, and the bleeding sap weight was significant positive correlated with bred era, the R4stage is the key to ensure root activity of soybean cultivation stage, through excellent cultivation measures to guarantee R4and any subsequent higher root activity is to ensure the core soybean yield cultivation technology means.
(3) The research results of the biomass and root biomass showed, that in V4period, the aboveground biomass of86years increased by10.81%, the average annual growth rate was0.13%; root biomass increased by28.10%, and the annual average growth rate was0.33%; individual biomass grew by11.68%, the average annual growth rate was0.14%; R2stage, the aboveground biomass increased by6.99%in the86years, the average annual growth rate was0.08%, root biomass increased by17.65%; root biomass average annual growth rate was0.21%; biomass of per plant increased by8.55%, the average annual growth rate was0.10%; R4stage, the aboveground biomass for86years increased by24.76%, the average annual growth rate was0.29%; root biomass increased by21.13%, the average annual growth rate was0.25%; biomass of per plant increased by24.21%, the average annual growth rate was0.28%; the aboveground biomass increased by10.51%in the86years at R6, the average annual growth rate was0.12%; root biomass increased by9.27%, the average annual growth rate was0.11%; biomass of per plant increased by10.05%, the average annual growth rate was0.12%. Therefore, root biomass, aboveground biomass and plant biomass in the R4stage showed a significant positive correlation or very significant positive correlation with bred era, the organ biomass of soybean varieties at V4, R2and R6were also positive correlated with bred era, but did not reach a significant level. This indicates that the ground and underground portion growth of soybean at R4played a crucial role in the production, the genetic improvement of soybean varieties significantly increased the plant organ biomass, these may be associated with the improvement of root biomass and root activity, especially root activity at R4, at the R4stage the xylem fluid weight and root activity were positively correlated with net photosynthetic rate, were also proved the relevance of the underground part and the ground photosynthesis. Therefore, we speculate that the root xylem fluid weight and root activity can be used as indicators of the net photosynthetic rate, and to provide theoretical basis for future breeding high photosynthetic soybean varieties.
(4) Root activity and root BSW of soybean varieties is closely related to the accumulation of dry matter. We also found that the aerial parts and the underground coordinated in improvement process, and the genetic improvement of soybean varieties was the result of coordination of the aboveground and underground organ. Correlation between root activity and root BSW and net photosynthetic rate of function leaf was the most significant at R4stage, and correlation between root activity and net photosynthetic rate did not reach significant level. The aerial parts and the underground coordinated better at R2and R4stage. At R6stage root activity significant declined, and root BSW quickly reduced, but net photosynthetic rate of leaves and biomass of underground parts decreased little. It showed that the decline of root activity was earlier than the decline of photosynthesis. So the reason of premature phenomenon in late aging with soybean cultivars may be resulted by the decline in photosynthesis in late aging with plants with the premature aging of roots. So we could delay aging of the root aimed to slow down the decline in photosynthesis in late growth aging, thereby to increase the yield.
该试验于2010年和2011年两年进行,选择1923年至2009年间育成的大豆品种进行研究,通过对不同年代育成大豆品种叶片光合速率、气孔导度、蒸腾速率、叶绿素和比叶重等光合指标的测定,以及对不同生育期根系伤流量和根系活力及地上和地下生物量的测定,进一步分析其之间的关系,研究结果表明：
(1)从1923年到2005年,82年来每公顷籽粒产量提高80.2%,平均每年增加0.98%。生育期82年来缩短了14.7天,平均每10年缩短1.83天,而产量随大豆品种熟期的缩短而呈增加的变化趋势；收获指数提高了58.03%,且与产量呈极显著正相关(r=0.8099**)；叶片的净光合速率增加了3.9196μmolCO2·m-1·s-1,提高了18.09%,平均每年提高0.22%,产量与叶片净光合速率呈极显著正相关(r=0.6102**)；叶片气孔导度提高了22.91%,平均每年提高0.27%,但产量与叶片气孔导度未达到显著正相关水平；叶片胞间二氧化碳浓度降低了8.3804μmolCO2·mol-1,平均每年降低0.1022μmolCO2·mol-1;蒸腾速率提高了57.29%,平均每年提高0.69%,产量随蒸腾速率的增加而增加,相关系数为0.6316**,而叶片水分利用效率随品种育成年代增加而显著减少,82年来减少了22.3%；叶片表观叶肉导度提高了49.54%,平均每年提高0.60%,产量随表观叶肉导度的增加而显著增加；叶片气孔限制值降低了8.25%,平均每年降低了0.10%,产量随叶片气孔限制值的增加而降低；叶片比叶重提高了91.82%,平均每年提高1.12%,产量随比叶重的增加而增加,相关系数为0.4464*；叶片叶绿素含量提高了17.41%,平均每年提高0.21%,产量随叶绿素的增加而增加；小叶面积降低了47.37%,平均每年降低0.58%。籽粒产量与Pn和Pn/Ci正相关达到极显著水平而叶片的水分利用效率随品种育成年代而显著降低与籽粒产量呈极显著负相关,这三个生理指标可以作为衡量籽粒产量的育种指标。
(2)从根系伤流液和根系活力看,在V4期,根系伤流量86年来增长了9.10%,年平均增长率为0.11%；根系活力增长了10.24%,年平均增长率为0.12%；在R2期,根系伤流量86年来增长了18.29%,年平均增长率为0.21%；根系活力增长了27.38%,年平均增长率为0.32%；在R4期,根系伤流量86年来增长了16.01%,年平均增长率为0.19%；根系活力增长了24.22%,年平均增长率为0.28%；在R6期,伤流液重量86年来增长了20.22%,年平均增长率为0.24%；根系活力增长了10.83%,年平均增长率为0.13%。由此可知,在R4期年增长率达到最大值,同时在R4期伤流液重量与育成年代呈显著正相关,R4期是大豆栽培中保证根系活力的关键阶段,通过优良的栽培措施保证R4期及以后较高的根系活力是保证大豆高产的核心栽培技术手段。
(3)从地上生物量和根系生物量看,在V4期,地上生物量86年来增长了10.81%,年平均增长率为0.13%；根系生物量增长了28.10%,年平均增长率为0.33%；单株生物量增长了11.68%,年平均增长率为0.14%；在R2期,地上部生物量86年来增长了6.99%,年平均增长率为0.08%；根生物量增长了17.65%,年平均增长率为0.21%；单株生物量增长了8.55%,年平均增长率为0.10%；在R4期,地上86年来增长了24.76%,年平均增长率为0.29%；根生物量增长了21.13%,年平均增长率为0.25%；单株生物量增长了24.21%,年平均增长率为0.28%；在R6期,地上部生物量86年来增长了10.51%,年平均增长率为0.12%；根生物量增长了9.27%,年平均增长率为0.11%；单株生物量增长了10.05%,年平均增长率为0.12%。由此可知,根生物量,地上部生物量和单株生物量在R4期与育成年代均呈显著正相关或极显著正相关,在V4、R2和R6期,大豆品种的器官生物量与育成年代虽也呈正相关,但均未达到显著水平。这表明,R4期大豆地上和地下部分的生长对产量起至关重要的作用,大豆品种的遗传改良使植株器官生物量得到显著增加,可能与根系生物量和根系活力的改善有关,特别是R4期的根系活力,在R4期,伤流液重量和根系活力与叶片净光合速率均呈显著正相关,也证明了地下部分与地上光合作用的相关性。因此我们推测根系伤流液重量和根系活力可以作为衡量叶片净光合速率的指标,为今后选育高光合大豆品种提供理论依据。
(4)大豆品种干物质的积累与根系活力、根系伤流液重量有着密切关系。同时我们也发现地上部分与地下根系在品种改良过程中,两者是协同进化的,大豆品种遗传改良是地上器官与地下器官协同进化的结果。在R4期,根系活力和根系伤流液重量和功能叶片的净光合速率关系最为显著,在R2期和R6期,根系活力和叶片净光合速率并未达到显著水平。R2期和R4期地上部分和地下部分有较好的协同关系,但在R6期,根系活力急剧下降,根系伤流液重量迅速减少,但地上部分器官生物量和叶片净光合速率下降不大,说明根系活力的下降先于光合作用的下降,因此大豆植株后期的早衰现象,可能是由于根系的提前衰老导致植株后期光合作用下降的原因。因此我们通过延缓根系的衰老,有可能延缓生育后期光合作用的下降,进而提高产量。
Many reports had showed about agronomic traits and physiological changes in the genetic improvement process of soybean cultivars, but about changes of roots in physiological functions have not been reported, especially about the relationship of root activity and aerial parts of organisms and photosynthesis. This study used soybean varieties bred in Jilin Province since1923to study the photosynthetic characteristics of soybean varieties and root BSW and the biomass of the aboveground, and analyze the relationship of photosynthetic characteristics and yield and the relationship between root BSW and root activity and biomass of aboveground. Aimed to clarify the evolution of the photosynthetic characteristics of leaves in the genetic improvement process of soybean cultivars, and reveal the relationship between the roots and aerial parts, to provide the theoretical basis for the the soybean varieties improvement in Jilin Province.
The experiment was conducted in2010and2011, and chose the soybean varieties bred in the1923-2009. Measured the photosynthetic rate, stomatal conductance, transpiration rate, chlorophyll and specific leaf weight of soybean cultivars, and measured the relationship of root BSW and root activity and biomass of aboveground to analyze their relationship further. The study results show that:
(1) From1923to2005, the grain yield per hectare in82years increased by80.2%, with an average increase by0.98%per year. The of growth period in82years was shortened by14.7days, with an average increase by1.83days every10year, while the yield were showed an increasing trend with shortening in growth period of soybean varieties; harvest index was raise by58.03%, and the harvest index was extremely significant correlated with yield (r=0.8099**); leaf net photosynthetic rate was increased by3.9196μmolCO2·m-1·s-1, and it was increased by18.09%, an average increase was by0.22%per year; the yield was highly significant positive correlated with net photosynthetic rate (r=0.6102**); leaf stomatal conductance was increased by22.91%, an average increase was by0.27%per year, but the correlation of yield and stomatal conductance did not reach significant level; leaf intercellular CO2concentration was decreased by8.3804μmolCO2·mol-1, an average decrease was by0.1022μmolCO2·mol-1per year; transpiration rate was increased by57.29%, an average increase was by0.69%per year, the yield was increased with the increase of transpiration rate, the correlation coefficient was0.6316, but the moisture use efficiency of leaves was decreased significantly along with the year of release, the decrease was by22.3%with the year of release in82years; the blades apparent mesophyll conductance was increased by49.54%, an average increase was by0.60%, the yield was significant increased with the increase in apparent mesophyll conductance; stomatal limitation in leaf was decreased by8.25%, and the average reduction was by0.10%per year, and the yield was decreased with the increase in stomatal limitation; the specific leaf weight was increased by91.82%, an average increase was by1.12%per year, the yield was increased with the increase in specific leaf weight, a correlation coefficient was0.4464; the chlorophyll content in leaf was increased by17.41%, with an average increase was by0.21%per year, the yield was increased with the increase in chlorophyll content; the area of leaflet was decreased by47.37%, with an average reduction was by0.58%. The yield was highly significant positive correlated with Pn, Pn/Ci, but the correlation between the water use efficiency in leaves and year of release was negative, and the water use efficiency in leaves was highly significant negative correlated with grain yield. In concluding, the three physiological indicators could be as indicators by measure of breeding of grain yield.
(2)The analysis on root bleeding sap and root vigor has showed that at V4stage, the root bleeding rate for86years increased by9.10%, the average annual growth rate was0.11%; the root activity grew by10.24%, the average annual growth rate was0.12%; at R2stage, the root bleeding rate increased by18.29%in the86years, the average annual growth rate was0.21%; the root activity grew by27.38%, the average annual growth rate was0.32%; at R4, root injury traffic in1986increased by16.01%, the average annual growth rate was0.19%; root activity grew by24.22%, the average annual growth rate was0.28%; R6of xylem sap weight increase by20.22%in the86years, the average annual growth rate was0.24%; growth of root activity was10.83%, and the average annual growth rate was0.13%. The result showed that at R4, the annual growth rate reached the maximum, and the bleeding sap weight was significant positive correlated with bred era, the R4stage is the key to ensure root activity of soybean cultivation stage, through excellent cultivation measures to guarantee R4and any subsequent higher root activity is to ensure the core soybean yield cultivation technology means.
(3) The research results of the biomass and root biomass showed, that in V4period, the aboveground biomass of86years increased by10.81%, the average annual growth rate was0.13%; root biomass increased by28.10%, and the annual average growth rate was0.33%; individual biomass grew by11.68%, the average annual growth rate was0.14%; R2stage, the aboveground biomass increased by6.99%in the86years, the average annual growth rate was0.08%, root biomass increased by17.65%; root biomass average annual growth rate was0.21%; biomass of per plant increased by8.55%, the average annual growth rate was0.10%; R4stage, the aboveground biomass for86years increased by24.76%, the average annual growth rate was0.29%; root biomass increased by21.13%, the average annual growth rate was0.25%; biomass of per plant increased by24.21%, the average annual growth rate was0.28%; the aboveground biomass increased by10.51%in the86years at R6, the average annual growth rate was0.12%; root biomass increased by9.27%, the average annual growth rate was0.11%; biomass of per plant increased by10.05%, the average annual growth rate was0.12%. Therefore, root biomass, aboveground biomass and plant biomass in the R4stage showed a significant positive correlation or very significant positive correlation with bred era, the organ biomass of soybean varieties at V4, R2and R6were also positive correlated with bred era, but did not reach a significant level. This indicates that the ground and underground portion growth of soybean at R4played a crucial role in the production, the genetic improvement of soybean varieties significantly increased the plant organ biomass, these may be associated with the improvement of root biomass and root activity, especially root activity at R4, at the R4stage the xylem fluid weight and root activity were positively correlated with net photosynthetic rate, were also proved the relevance of the underground part and the ground photosynthesis. Therefore, we speculate that the root xylem fluid weight and root activity can be used as indicators of the net photosynthetic rate, and to provide theoretical basis for future breeding high photosynthetic soybean varieties.
(4) Root activity and root BSW of soybean varieties is closely related to the accumulation of dry matter. We also found that the aerial parts and the underground coordinated in improvement process, and the genetic improvement of soybean varieties was the result of coordination of the aboveground and underground organ. Correlation between root activity and root BSW and net photosynthetic rate of function leaf was the most significant at R4stage, and correlation between root activity and net photosynthetic rate did not reach significant level. The aerial parts and the underground coordinated better at R2and R4stage. At R6stage root activity significant declined, and root BSW quickly reduced, but net photosynthetic rate of leaves and biomass of underground parts decreased little. It showed that the decline of root activity was earlier than the decline of photosynthesis. So the reason of premature phenomenon in late aging with soybean cultivars may be resulted by the decline in photosynthesis in late aging with plants with the premature aging of roots. So we could delay aging of the root aimed to slow down the decline in photosynthesis in late growth aging, thereby to increase the yield.
引文
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