光强对花生光合特性、产量和品质的影响及生长模型研究
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摘要
本研究于2007年和2008年在山东农业大学农学实验站进行。针对花生与玉米、小麦等作物间作、套种造成遮光问题,选用大花生品种丰花1号和小花生品种丰花2号做为供试材料,在花生苗期、结荚期和饱果期分别用不同透光率的遮阳网进行遮光处理模拟弱光条件,设置自然光(CK)、遮光27%、遮光43%和遮光77%四种光强水平,研究了光强处理对花生光合特性、光系统Ⅱ光能分配、RuBPCase活性、植株形态建成、物质生产和产量以及品质的影响,明确了不同时期干物质积累及光能利用率与光合有效辐射的关系,以此为基础采用2种方法建立了花生干物质生产动态模型和花生产量形成模拟模型,并对各模型的模拟精度进行了比较。主要研究结果如下:
1光强对花生叶片光合荧光特性的影响
苗期遮光处理,随处理光强的减弱叶绿素(Chl(a+b))含量显著增加,PSⅡ的实际光化学效率(ФPSⅡ)和最大光化学效率(Fv/Fm)升高;叶绿素a(Chla)与叶绿素b(Chlb)的比值(Chla/b)、净光合速率(Pn)降低。遮光处理结束,各处理均在高光强1200μmol·m-2·s-1下测定时,Pn、气孔导度(Gs)随处理光强的减弱而下降,细胞间隙CO2浓度(Ci)升高;在低光强276μmol·m-2·s-1下测定时,遮光处理的Pn显著高于自然光下生长的,Gs和Ci下降;低光强与高光强下测定的Pn的比值随处理光强的减弱显著升高。遮光处理结束后均恢复到自然光强下,Pn、ФPSⅡ和Fv/Fm先迅速下降,3-5d后逐渐回升;恢复15d时,仅遮光27%处理的各参数能恢复到对照水平。丰花1号各处理叶绿素含量、Pn、ФPSⅡ均比丰花2号相同处理的高。结荚期遮光处理花生叶的Pn变化规律与苗期的类似。饱果期遮光处理的在处理光强下和低光强276μmol·m-2·s-1光强下测定时Pn变化规律与苗期处理的相似,均在高光强1200μmol·m-2·s-1下测定时Pn随处理光强的减弱而升高。遮光处理降低了花生的光合能力,提高了其利用弱光的能力,降低了其利用强光的能力;并且花生对弱光有一定有适应性,轻度遮光处理的在自然光下生长一段时间可以较快的恢复。
2光强对光合日变化及光合曲线参数的影响
花生叶片Pn的日变化都呈单峰曲线,遮光处理对Pn日变化的趋势没有影响,但降低了Pn的峰值,提高了花生下午弱光时的光合速率。苗期在遮光条件下生长的植株光合作用光补偿点和光饱和点比自然光强下生长的植株显著降低,表观量子效率显著升高,光强越弱变幅越大。遮光27%处理的光饱和点与自然光强下的差异不显著,遮光43%和77%处理的光饱和点分别比自然光强下的降低14%、29%,差异显著。自然光、遮光27%、43%和77%处理的光合作用光补偿点分别为:52.7、45.9、21.3、9.6μmol·m-2·s-1,表观量子效率为:0.0269、0.0317、0.0337、0.0317μmol·mol-1。弱光处理后,叶片CO2补偿点、CO2饱和点降低,遮光43%和77%处理的与自然光下生长的差异显著。
3光强对光合酶及抗氧化酶活性的影响
苗期遮光处理的花生叶片RuBPCase活性都显著地低于自然光下生长的,且随处理光强的减弱而显著降低,遮光27%、43%和77%处理分别比对照降低21.7%、45.9%和81.9%。苗期遮光处理结束,均在自然光下生长1d,消除了即时光强的刺激,遮光处理的RUBPCase有所升高,但仍随处理光强的减弱而显著降低,表现出RUBPCase的潜力差异,苗期遮光可能损害了叶片的RuBPCase功能。PEPCase活性变化规律与RUBPCase活性基本一致,遮光处理的都显著地低于自然光下生长的,且随处理光强的减弱而降低,但降低的幅度比RUBPCase活性的小。说明即时光强和较长时间的遮光处理均会显著影响RuBPCase和PEPCase的活性。
苗期遮光处理,随处理光强的减弱,花生叶片的SOD、POD活性显著升高,CAT活性显著降低;处理结束后在自然光下生长15d时,SOD、POD活性仍随处理光强的减弱而升高,但与对照差异减小,CAT活性仍比自然光下生长的低。结荚期和饱果期遮光处理的SOD活性随处理光强的减弱而下降,POD和CAT活性上升。结荚期遮光处理结束在自然光下生长15d时,各处理SOD活性均明显低于对照,POD和CAT活性高于对照,但处理间差异不显著。
4光强对花生植株营养生长的影响
苗期遮光处理对花生植株性状影响最大,结荚期处理的次之,饱果期处理的影响最小。苗期遮光处理促进了花生主茎和侧枝的伸长,但明显减少了花生植株的分枝数。3个时期遮光处理对叶面积系数均有很大影响,其中苗期遮光43%和77%处理主要是降低了叶面积的增长速度,从而降低了最大叶面积系数;结荚期遮光处理则是减缓或是阻止了花生生长后期叶面积系数的提高,从而使最大叶面积系数变小,出现的时间比自然光下生长的早,同时减缓叶面积系数下降的速度;饱果期遮光处理则是加快了叶面积系数下降的速度。3个时期遮光处理植株生物产量均随处理光强的减弱显著降低,其中以结荚期遮光处理的降幅最大,饱果期遮光处理的次之,苗期遮光处理的影响最小。
5光强对荚果产量和籽仁品质的影响
遮光处理均造成减产,苗期遮光27%和43%处理的减产幅度较小与对照差异不显著;结荚期和饱果期遮光处理的减产严重差异显著,减产原因是降低了单株结果数和荚果的成熟饱满度。苗期遮光处理对花生产品质量影响较小,结荚期和饱果期遮光处理影响较大,出口果和仁出成率均降低,籽仁含油量分别降低0.4～2.0、0.6～3.2、1.6～6.0个百分点,蛋白质和可溶性糖含量小幅增加。籽仁典型样品(成熟饱满的花生籽仁)含油量比随机样品(具有经济价值的籽仁,包括成熟饱满的和秕的)分别高0.7～2.6、0.7～3.4、0.7～4.9个百分点。籽仁营养成分含量和成熟饱满度极显著相关,说明通过栽培措施提高成熟饱满度的途径改善花生产品质量有很大潜力。
6干物质积累、光能利用率与光合有效辐射的关系
系统研究了不同生育时期光能利用率、干物质积累与光合有效辐射的关系。苗期单株干物质增长速率相应与光合有效辐射量的变化符合双曲线,结荚期的符合指数曲线,饱果期的符合S形曲线,3个时期的相对增长速率相应与光合有效辐射量的变化均符合指数曲线;结荚期和饱果期荚果干物质增长速率和相对增长速率与光合有效辐射均符合指数曲线。为建立以光合有效辐射为驱动变量的干物质积累模型提供了基础。以光合有效辐射为基数计算了花生的光能利用率,全生育期平均为5.19%,苗期平均为2.33%,花针期平均为7.33%,结荚期平均为8.87%,饱果期平均为7.72%。苗期光能利用率与光合有效辐射负相关,结荚期和饱果期是正相关;光能利用率与叶面积系数呈正相关关系。
7花生干物质生产与产量形成模拟模型
在试验研究的基础上,建立了花生干物质生产模型,然后以此为基础建立花生产量形成模拟模型,并进行了比较。第一种方法是根据“物质—能量转化—能量平衡”理论及作物生理学的基本原理,建立了花生群体干物质生产动态模型。模型考虑了群体叶面积动态、冠层光能分布及光合作用等主要生理过程。该模型的群体干物质积累实测值与预测值的决定系数为0.9505,均方差根为16.76%,以此为基础进行的产量预测检验结果为:R=0.9236,RMSE=17.58%。二是以光合有效辐射Q与叶面积系数L作为驱动变量,借鉴CERES中的干物质生产模型Wi=a*Qi*(1-e-k*Li),按出苗—结荚期,结荚期—成熟收获两个阶段进行模拟,干物质积累实测值与预测值的决定系数为0.9967,均方差根为7.18%,以此为基础进行的产量预测检验结果为:R=0.9758,RMSE=9.88%。两种模型相比,第一种方法建立的模型机理性较强,CERES模型有更好的模拟精度。
The research was carried out at agronomy experimental station of Shandong agricultural university in 2007 and 2008. Shading experiments were designed according to low light intensity problem caused by intercropping system of peanut with maize or wheat et al. Fenghua1 and Fenghua2 were adopted in the experiment of four light intensity levels (CK in which the plants were grown under natural light, 27% shading, 43% shading and 77% shading), which carried out at three growth stage using black sunshade net. And the three stages were seedling phase, pod-setting phase and pod-maturing phase. The aim of this study was to investigate the effects of low light on the photosynthetic characteristics, luminous energy distribute of PSⅡ, RuBPCase activitiy, plant morphogenesis, dry matter production, pod yield and quality of peanut at different stages, defined the relationship between dry matter accumulation and PAR, relationship of efficiency for solar energy utilization and PAR, and then set up the dry matter production and yield formation model on the basis of the experimental data. The main results were as follows.
1 Effect of light intensity on photosynthetic and fluorescence characteristics of peanut leaves
Shading treatment at seedling phase could increase content of chlorophyll(a+b) (Chl(a+b)), enhance the actual photochemical efficiency of PSⅡ(ФPSⅡ) and optimal photochemical efficiency of PSⅡin the dark(Fv/Fm) significantly along with the weakening of treatment light intensity. While could depress the ratio of chlorophyll a to chlorophyll b (Chla/b) and photosynthetic rate (Pn). When the treatment ended, Pn, stomatal conductance (Gs) decreased and intercellular CO2 concentration (Ci) increased with the weakening of treatment light intensity when measured under high light intensity (1200μmol·m-2·s-1). And Pn raised, Gs and Ci reduced when measured under low light intensity (276μmol·m-2·s-1). The ratio of Pn measured in low light to high light had significant positive-correlation with treatment light intensity. When the shading treatment ended and recovered to natural light, Pn,ФPSⅡand Fv/Fm were immediately decreased and then ascended gradually after 3-5 days. The recovery extent and shading degree were negative correlation. After recovery of 15 days, Pn,ФPSⅡand Fv/Fm of 27% shading treatment could recover to the CK level. Content of Chl(a+b), Pn andФPSⅡof Fenghua1 were higher than those of Fenghua2 at the same treatment. So the ability of using low light was improved and the ability of using high light was decreased. The shading treatment at pod-setting phase had similar rule. Pn of shading treatments at pod-maturing phase had similar variation law with seedling treatment when measured under treatment light intensity and low light intensity (276μmol·m-2·s-1), while had opposite law when measured under high light intensity (1200μmol·m-2·s-1). So shading treatment depressed the photosynthetic capacity of peanut leaves, improved the ability of using low light, and decreased the ability of using high light. Peanut has adaptability to weak light, and peanut under light shading treatment resume growing in a period when growth in natural light.
2 Effect of light intensity on diurnal variation of photosynthesis and photosynthetic curve parameter of peanut leaves
The diurnal variation of net photosynthetic rate was a single peak curve. Shading treatment has no effect on the tendency of change, but depressed the peak of the curve and increased Pn in weak light in the afternoon. In seedling shading treatment, light compensation point and light saturation point of peanut which cultured under treatment light intensity conditions were lower than that cultured under natural light, while the apparent quantum yield were higher. The weaker the light intensity was, the larger amplitudes were. Compared with peanut cultured under nature light, light saturation point of peanut of 27% shading had not significant difference, while the light saturation point of peanut of 43% shading and 77% shading percentage fell by 14%, 29% respectively and were remarkable difference. The light compensation point of nature light, 27% shading, 43% shading and 77% shading were respectively 52.7, 45.9, 21.3, 9.6μmol·m-2·s-1, and the apparent quantum yield of the four treatment were respectively 0.0269, 0.0317, 0.0337, 0.0317μmol·mol-1. Shading treatment depressed the CO2 compensation point and CO2 saturation point, and the two indexes of 43% shading and 77% shading treatment had significant difference from the contrast.
3 Effect of light intensity on photosynthetic enzymatic and antioxidant enzyme activity of peanut leaves
RuBPCase activity of peanut leaves which cultured under low light ware significantly depressed according to the weakening of treatment light intensity in seedling phase. RuBPCase activity of peanut cultured under 27% shading, 43% shading and 77% shading treatment condition were decreased by 21.7%, 45.9% and 81.9% respectively compared with those cultured under natural light. When the seedling shading treatment ended, all the peanut plant were made growth in natural light for one day in order to eliminate the influence of immediate light intensity effect. RuBPCase activity increased slightly but also significantly depressed according to the weakening of treatment light intensity. The later data showed the difference of RuBPCase activity potential, that is to say seedling shading treatment was likely to damage the function of RuBPCase. PEPCase activity had the similar variation law to RuBPCase activity, while had lower amplitudes. The result indicated that immediate light intensity and long time shading treatment can all influence RuBPCase and PEPCase activity.
SOD, POD activity of peanut leaves cultured under low light were significantly increased according to the weakening of treatment light intensity at seedling phase, while CAT activity decreased significantly. After 15 days’growth under natural light when the treatment ended, SOD, POD, CAT activity all showed similar rules, while the disparity between the CK level minished. SOD activity depressed as the weakening of treatment light intensity in pod-setting phase and pod-maturing phase treatment, while POD and CAT activity increased. The SOD activity was significantly lower than CK level after 15 days’growth under natural light when the pod-setting shading treatment ended, while POD and CAT activity were higher than CK level but had not significantly difference.
4 Effect of light intensity on vegetative growth of peanut
Shading treatment at seedling phase had the most influential effect on peanut plant characters; shading at pod-setting phase took the second place; while shading at pod-maturing phase had the least influence. Shading treatment at seedling phase promoted the elongation of main stem and lateral branch, but reduced the branch number of the plant. Shading at the three phase all have great influence on leaf area index; among them 43% shading and 77% shading at seedling phase decreased leaf area index mainly because depressed the growth rate; shading at pod-setting phase mainly reduced the growth rate of leaf area, and depressed the descending rate at anaphase of peanut growth; shading at pod-maturing phase mainly quickened the degressive velocity of leaf area. Plant biolobical yield all significantly reduced at the three phase shading treatment, among them shading treatment at pod-setting had the extremum decreasing amplitude, shading treatment at pod-maturing phase took the second place, while shading at seedling phase had the least influence.
5 Effect of light intensity on pod yield and kernel quality of peanut
Yield reduction occurred under all treatments. There were little reduction under 27% and 43% shading at seedling phase, while the maximal yield reduction occurred under shading at pod-setting phase. Decrease of pods per plant and plumpness were main reason of reduction of yield. Shading at seedling phase had little influence on quality, while had great influence at the other two phases. The yield rate of export pod and kernel were all decreased. Decreasing ranges of oil content were 0.4～2.0, 0.6～3.2 and 1.6～6.0 percentage point at the three phases respectively, while protein and total soluble sugar content increased by a small margin. The oil content of typical samples was 0.7～2.6, 0.7～3.4 and 0.7～4.9 percentage point higher than random samples at the three phases respectively. There were significant correlation between nutrition content and plumpness. So it was very potential in improving the quality of peanut production by way of increasing plumpness with cultivation measures.
6 Relationships between dry matter accumulation and PAR, and between efficiency for solar energy utilization and PAR were specified
The relationships between dry matter accumulation and PAR and the relationship between efficiency for solar energy utilization and PRA were studied systemically. And it showed dry matter accumulation rate and PAR followed the logarithm curve at seedling phase, exponential curve at pod-setting phase and S shape curve at pod-maturing phase. Relative rate of dry matter accumulation of the three phases were all followed exponential curve. And the two rates of dry matter accumulate in pod were all followed exponential curve at pod-setting phase and pod-maturing phase. That’s to say there’s significant correlation between dry matter accumulation and PAR, and it will be the basic to make dry matter accumulation model. Efficiency for solar energy utilization was calculated, and the results as followed: 5.19% in the whole growth period, 2.33% at seedling phase, 7.33% at pod-pin phase, 8.87% at pod-setting phase and 7.72% in pod-maturing phase. There were negative correlation between efficiency for solar energy utilization and PAR at seedling phase, and positive correlation at pod-setting phase and pod-maturing phase. There were positive correlation between efficiency for solar energy utilization and LAI.
7 Simulation model of peanut population dry mater accumulation and yield formation were set up
Peanut population dry matter production model and yield formation model were founded by adopting two kinds of method based on the theories of peanut growth physiology. The first kind of model considered the physiological processes of leaf area dynamic, coronal luminous energy allocation, photosynthesis and so on based on the“material---energy transform--- energy balancing”theory and crop physiology fundamental principle. The test result of population dry matter production model is: R=0.9505, RMSE=16.76%; and test result of yield formation model is: R=0.9236, RMSE=17.58%. That’s the first peanut model found by this way. In the second kinds, solar radiation and leaf area index had been taken into account in dry matter model Wi=a*Qi*(1-e-k*Li) as CERES used for reference. According as the growth center is different in every period, the entire phenology course is detached to two phases, seedling to Pod-setting and Pod-setting to Pod-maturing. The test result of population dry matter production model is: R=0.9967, RMSE=7.18%; and test result of yield formation model is: R=0.9758, RMSE=9.88%. As compared, the first kind of model was much more mechanism, and the CERES model was batter in simulating precision.
1光强对花生叶片光合荧光特性的影响
苗期遮光处理,随处理光强的减弱叶绿素(Chl(a+b))含量显著增加,PSⅡ的实际光化学效率(ФPSⅡ)和最大光化学效率(Fv/Fm)升高;叶绿素a(Chla)与叶绿素b(Chlb)的比值(Chla/b)、净光合速率(Pn)降低。遮光处理结束,各处理均在高光强1200μmol·m-2·s-1下测定时,Pn、气孔导度(Gs)随处理光强的减弱而下降,细胞间隙CO2浓度(Ci)升高;在低光强276μmol·m-2·s-1下测定时,遮光处理的Pn显著高于自然光下生长的,Gs和Ci下降;低光强与高光强下测定的Pn的比值随处理光强的减弱显著升高。遮光处理结束后均恢复到自然光强下,Pn、ФPSⅡ和Fv/Fm先迅速下降,3-5d后逐渐回升;恢复15d时,仅遮光27%处理的各参数能恢复到对照水平。丰花1号各处理叶绿素含量、Pn、ФPSⅡ均比丰花2号相同处理的高。结荚期遮光处理花生叶的Pn变化规律与苗期的类似。饱果期遮光处理的在处理光强下和低光强276μmol·m-2·s-1光强下测定时Pn变化规律与苗期处理的相似,均在高光强1200μmol·m-2·s-1下测定时Pn随处理光强的减弱而升高。遮光处理降低了花生的光合能力,提高了其利用弱光的能力,降低了其利用强光的能力;并且花生对弱光有一定有适应性,轻度遮光处理的在自然光下生长一段时间可以较快的恢复。
2光强对光合日变化及光合曲线参数的影响
花生叶片Pn的日变化都呈单峰曲线,遮光处理对Pn日变化的趋势没有影响,但降低了Pn的峰值,提高了花生下午弱光时的光合速率。苗期在遮光条件下生长的植株光合作用光补偿点和光饱和点比自然光强下生长的植株显著降低,表观量子效率显著升高,光强越弱变幅越大。遮光27%处理的光饱和点与自然光强下的差异不显著,遮光43%和77%处理的光饱和点分别比自然光强下的降低14%、29%,差异显著。自然光、遮光27%、43%和77%处理的光合作用光补偿点分别为:52.7、45.9、21.3、9.6μmol·m-2·s-1,表观量子效率为:0.0269、0.0317、0.0337、0.0317μmol·mol-1。弱光处理后,叶片CO2补偿点、CO2饱和点降低,遮光43%和77%处理的与自然光下生长的差异显著。
3光强对光合酶及抗氧化酶活性的影响
苗期遮光处理的花生叶片RuBPCase活性都显著地低于自然光下生长的,且随处理光强的减弱而显著降低,遮光27%、43%和77%处理分别比对照降低21.7%、45.9%和81.9%。苗期遮光处理结束,均在自然光下生长1d,消除了即时光强的刺激,遮光处理的RUBPCase有所升高,但仍随处理光强的减弱而显著降低,表现出RUBPCase的潜力差异,苗期遮光可能损害了叶片的RuBPCase功能。PEPCase活性变化规律与RUBPCase活性基本一致,遮光处理的都显著地低于自然光下生长的,且随处理光强的减弱而降低,但降低的幅度比RUBPCase活性的小。说明即时光强和较长时间的遮光处理均会显著影响RuBPCase和PEPCase的活性。
苗期遮光处理,随处理光强的减弱,花生叶片的SOD、POD活性显著升高,CAT活性显著降低;处理结束后在自然光下生长15d时,SOD、POD活性仍随处理光强的减弱而升高,但与对照差异减小,CAT活性仍比自然光下生长的低。结荚期和饱果期遮光处理的SOD活性随处理光强的减弱而下降,POD和CAT活性上升。结荚期遮光处理结束在自然光下生长15d时,各处理SOD活性均明显低于对照,POD和CAT活性高于对照,但处理间差异不显著。
4光强对花生植株营养生长的影响
苗期遮光处理对花生植株性状影响最大,结荚期处理的次之,饱果期处理的影响最小。苗期遮光处理促进了花生主茎和侧枝的伸长,但明显减少了花生植株的分枝数。3个时期遮光处理对叶面积系数均有很大影响,其中苗期遮光43%和77%处理主要是降低了叶面积的增长速度,从而降低了最大叶面积系数;结荚期遮光处理则是减缓或是阻止了花生生长后期叶面积系数的提高,从而使最大叶面积系数变小,出现的时间比自然光下生长的早,同时减缓叶面积系数下降的速度;饱果期遮光处理则是加快了叶面积系数下降的速度。3个时期遮光处理植株生物产量均随处理光强的减弱显著降低,其中以结荚期遮光处理的降幅最大,饱果期遮光处理的次之,苗期遮光处理的影响最小。
5光强对荚果产量和籽仁品质的影响
遮光处理均造成减产,苗期遮光27%和43%处理的减产幅度较小与对照差异不显著;结荚期和饱果期遮光处理的减产严重差异显著,减产原因是降低了单株结果数和荚果的成熟饱满度。苗期遮光处理对花生产品质量影响较小,结荚期和饱果期遮光处理影响较大,出口果和仁出成率均降低,籽仁含油量分别降低0.4～2.0、0.6～3.2、1.6～6.0个百分点,蛋白质和可溶性糖含量小幅增加。籽仁典型样品(成熟饱满的花生籽仁)含油量比随机样品(具有经济价值的籽仁,包括成熟饱满的和秕的)分别高0.7～2.6、0.7～3.4、0.7～4.9个百分点。籽仁营养成分含量和成熟饱满度极显著相关,说明通过栽培措施提高成熟饱满度的途径改善花生产品质量有很大潜力。
6干物质积累、光能利用率与光合有效辐射的关系
系统研究了不同生育时期光能利用率、干物质积累与光合有效辐射的关系。苗期单株干物质增长速率相应与光合有效辐射量的变化符合双曲线,结荚期的符合指数曲线,饱果期的符合S形曲线,3个时期的相对增长速率相应与光合有效辐射量的变化均符合指数曲线;结荚期和饱果期荚果干物质增长速率和相对增长速率与光合有效辐射均符合指数曲线。为建立以光合有效辐射为驱动变量的干物质积累模型提供了基础。以光合有效辐射为基数计算了花生的光能利用率,全生育期平均为5.19%,苗期平均为2.33%,花针期平均为7.33%,结荚期平均为8.87%,饱果期平均为7.72%。苗期光能利用率与光合有效辐射负相关,结荚期和饱果期是正相关;光能利用率与叶面积系数呈正相关关系。
7花生干物质生产与产量形成模拟模型
在试验研究的基础上,建立了花生干物质生产模型,然后以此为基础建立花生产量形成模拟模型,并进行了比较。第一种方法是根据“物质—能量转化—能量平衡”理论及作物生理学的基本原理,建立了花生群体干物质生产动态模型。模型考虑了群体叶面积动态、冠层光能分布及光合作用等主要生理过程。该模型的群体干物质积累实测值与预测值的决定系数为0.9505,均方差根为16.76%,以此为基础进行的产量预测检验结果为:R=0.9236,RMSE=17.58%。二是以光合有效辐射Q与叶面积系数L作为驱动变量,借鉴CERES中的干物质生产模型Wi=a*Qi*(1-e-k*Li),按出苗—结荚期,结荚期—成熟收获两个阶段进行模拟,干物质积累实测值与预测值的决定系数为0.9967,均方差根为7.18%,以此为基础进行的产量预测检验结果为:R=0.9758,RMSE=9.88%。两种模型相比,第一种方法建立的模型机理性较强,CERES模型有更好的模拟精度。
The research was carried out at agronomy experimental station of Shandong agricultural university in 2007 and 2008. Shading experiments were designed according to low light intensity problem caused by intercropping system of peanut with maize or wheat et al. Fenghua1 and Fenghua2 were adopted in the experiment of four light intensity levels (CK in which the plants were grown under natural light, 27% shading, 43% shading and 77% shading), which carried out at three growth stage using black sunshade net. And the three stages were seedling phase, pod-setting phase and pod-maturing phase. The aim of this study was to investigate the effects of low light on the photosynthetic characteristics, luminous energy distribute of PSⅡ, RuBPCase activitiy, plant morphogenesis, dry matter production, pod yield and quality of peanut at different stages, defined the relationship between dry matter accumulation and PAR, relationship of efficiency for solar energy utilization and PAR, and then set up the dry matter production and yield formation model on the basis of the experimental data. The main results were as follows.
1 Effect of light intensity on photosynthetic and fluorescence characteristics of peanut leaves
Shading treatment at seedling phase could increase content of chlorophyll(a+b) (Chl(a+b)), enhance the actual photochemical efficiency of PSⅡ(ФPSⅡ) and optimal photochemical efficiency of PSⅡin the dark(Fv/Fm) significantly along with the weakening of treatment light intensity. While could depress the ratio of chlorophyll a to chlorophyll b (Chla/b) and photosynthetic rate (Pn). When the treatment ended, Pn, stomatal conductance (Gs) decreased and intercellular CO2 concentration (Ci) increased with the weakening of treatment light intensity when measured under high light intensity (1200μmol·m-2·s-1). And Pn raised, Gs and Ci reduced when measured under low light intensity (276μmol·m-2·s-1). The ratio of Pn measured in low light to high light had significant positive-correlation with treatment light intensity. When the shading treatment ended and recovered to natural light, Pn,ФPSⅡand Fv/Fm were immediately decreased and then ascended gradually after 3-5 days. The recovery extent and shading degree were negative correlation. After recovery of 15 days, Pn,ФPSⅡand Fv/Fm of 27% shading treatment could recover to the CK level. Content of Chl(a+b), Pn andФPSⅡof Fenghua1 were higher than those of Fenghua2 at the same treatment. So the ability of using low light was improved and the ability of using high light was decreased. The shading treatment at pod-setting phase had similar rule. Pn of shading treatments at pod-maturing phase had similar variation law with seedling treatment when measured under treatment light intensity and low light intensity (276μmol·m-2·s-1), while had opposite law when measured under high light intensity (1200μmol·m-2·s-1). So shading treatment depressed the photosynthetic capacity of peanut leaves, improved the ability of using low light, and decreased the ability of using high light. Peanut has adaptability to weak light, and peanut under light shading treatment resume growing in a period when growth in natural light.
2 Effect of light intensity on diurnal variation of photosynthesis and photosynthetic curve parameter of peanut leaves
The diurnal variation of net photosynthetic rate was a single peak curve. Shading treatment has no effect on the tendency of change, but depressed the peak of the curve and increased Pn in weak light in the afternoon. In seedling shading treatment, light compensation point and light saturation point of peanut which cultured under treatment light intensity conditions were lower than that cultured under natural light, while the apparent quantum yield were higher. The weaker the light intensity was, the larger amplitudes were. Compared with peanut cultured under nature light, light saturation point of peanut of 27% shading had not significant difference, while the light saturation point of peanut of 43% shading and 77% shading percentage fell by 14%, 29% respectively and were remarkable difference. The light compensation point of nature light, 27% shading, 43% shading and 77% shading were respectively 52.7, 45.9, 21.3, 9.6μmol·m-2·s-1, and the apparent quantum yield of the four treatment were respectively 0.0269, 0.0317, 0.0337, 0.0317μmol·mol-1. Shading treatment depressed the CO2 compensation point and CO2 saturation point, and the two indexes of 43% shading and 77% shading treatment had significant difference from the contrast.
3 Effect of light intensity on photosynthetic enzymatic and antioxidant enzyme activity of peanut leaves
RuBPCase activity of peanut leaves which cultured under low light ware significantly depressed according to the weakening of treatment light intensity in seedling phase. RuBPCase activity of peanut cultured under 27% shading, 43% shading and 77% shading treatment condition were decreased by 21.7%, 45.9% and 81.9% respectively compared with those cultured under natural light. When the seedling shading treatment ended, all the peanut plant were made growth in natural light for one day in order to eliminate the influence of immediate light intensity effect. RuBPCase activity increased slightly but also significantly depressed according to the weakening of treatment light intensity. The later data showed the difference of RuBPCase activity potential, that is to say seedling shading treatment was likely to damage the function of RuBPCase. PEPCase activity had the similar variation law to RuBPCase activity, while had lower amplitudes. The result indicated that immediate light intensity and long time shading treatment can all influence RuBPCase and PEPCase activity.
SOD, POD activity of peanut leaves cultured under low light were significantly increased according to the weakening of treatment light intensity at seedling phase, while CAT activity decreased significantly. After 15 days’growth under natural light when the treatment ended, SOD, POD, CAT activity all showed similar rules, while the disparity between the CK level minished. SOD activity depressed as the weakening of treatment light intensity in pod-setting phase and pod-maturing phase treatment, while POD and CAT activity increased. The SOD activity was significantly lower than CK level after 15 days’growth under natural light when the pod-setting shading treatment ended, while POD and CAT activity were higher than CK level but had not significantly difference.
4 Effect of light intensity on vegetative growth of peanut
Shading treatment at seedling phase had the most influential effect on peanut plant characters; shading at pod-setting phase took the second place; while shading at pod-maturing phase had the least influence. Shading treatment at seedling phase promoted the elongation of main stem and lateral branch, but reduced the branch number of the plant. Shading at the three phase all have great influence on leaf area index; among them 43% shading and 77% shading at seedling phase decreased leaf area index mainly because depressed the growth rate; shading at pod-setting phase mainly reduced the growth rate of leaf area, and depressed the descending rate at anaphase of peanut growth; shading at pod-maturing phase mainly quickened the degressive velocity of leaf area. Plant biolobical yield all significantly reduced at the three phase shading treatment, among them shading treatment at pod-setting had the extremum decreasing amplitude, shading treatment at pod-maturing phase took the second place, while shading at seedling phase had the least influence.
5 Effect of light intensity on pod yield and kernel quality of peanut
Yield reduction occurred under all treatments. There were little reduction under 27% and 43% shading at seedling phase, while the maximal yield reduction occurred under shading at pod-setting phase. Decrease of pods per plant and plumpness were main reason of reduction of yield. Shading at seedling phase had little influence on quality, while had great influence at the other two phases. The yield rate of export pod and kernel were all decreased. Decreasing ranges of oil content were 0.4～2.0, 0.6～3.2 and 1.6～6.0 percentage point at the three phases respectively, while protein and total soluble sugar content increased by a small margin. The oil content of typical samples was 0.7～2.6, 0.7～3.4 and 0.7～4.9 percentage point higher than random samples at the three phases respectively. There were significant correlation between nutrition content and plumpness. So it was very potential in improving the quality of peanut production by way of increasing plumpness with cultivation measures.
6 Relationships between dry matter accumulation and PAR, and between efficiency for solar energy utilization and PAR were specified
The relationships between dry matter accumulation and PAR and the relationship between efficiency for solar energy utilization and PRA were studied systemically. And it showed dry matter accumulation rate and PAR followed the logarithm curve at seedling phase, exponential curve at pod-setting phase and S shape curve at pod-maturing phase. Relative rate of dry matter accumulation of the three phases were all followed exponential curve. And the two rates of dry matter accumulate in pod were all followed exponential curve at pod-setting phase and pod-maturing phase. That’s to say there’s significant correlation between dry matter accumulation and PAR, and it will be the basic to make dry matter accumulation model. Efficiency for solar energy utilization was calculated, and the results as followed: 5.19% in the whole growth period, 2.33% at seedling phase, 7.33% at pod-pin phase, 8.87% at pod-setting phase and 7.72% in pod-maturing phase. There were negative correlation between efficiency for solar energy utilization and PAR at seedling phase, and positive correlation at pod-setting phase and pod-maturing phase. There were positive correlation between efficiency for solar energy utilization and LAI.
7 Simulation model of peanut population dry mater accumulation and yield formation were set up
Peanut population dry matter production model and yield formation model were founded by adopting two kinds of method based on the theories of peanut growth physiology. The first kind of model considered the physiological processes of leaf area dynamic, coronal luminous energy allocation, photosynthesis and so on based on the“material---energy transform--- energy balancing”theory and crop physiology fundamental principle. The test result of population dry matter production model is: R=0.9505, RMSE=16.76%; and test result of yield formation model is: R=0.9236, RMSE=17.58%. That’s the first peanut model found by this way. In the second kinds, solar radiation and leaf area index had been taken into account in dry matter model Wi=a*Qi*(1-e-k*Li) as CERES used for reference. According as the growth center is different in every period, the entire phenology course is detached to two phases, seedling to Pod-setting and Pod-setting to Pod-maturing. The test result of population dry matter production model is: R=0.9967, RMSE=7.18%; and test result of yield formation model is: R=0.9758, RMSE=9.88%. As compared, the first kind of model was much more mechanism, and the CERES model was batter in simulating precision.
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