‘寒富’苹果光合生理及碳素物质代谢研究
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摘要
‘寒富'苹果的育成在扩大大苹果栽培区域,构建大苹果新的优势产业带中发挥了重要作用,但其相关生理基础和技术措施的作用机制研究十分缺乏,尤其是冷凉气候条件对优质大苹果植株光合特性及相关生理基础影响的研究尚未见报道。本试验以‘寒富'苹果及其亲本‘东光'和‘富士'为试材,采用光合气体交换及叶绿素荧光分析技术等手段,结合组织解剖特征分析和生理生化指标测定,对‘寒富'苹果的光合生理及碳素物质代谢进行初步评价,通过人为创造各种胁迫环境,研究不同土壤管理制度条件下‘寒富'苹果各器官的光合生理特性及碳素物质代谢基础,探讨‘寒富'苹果的光合生理及碳素物质代谢机制,揭示冷凉气候区‘寒富'苹果多抗性表达的生理基础,以期为冷凉气候区‘寒富'苹果的优质栽培管理提供理论依据。主要研究结果如下:
1‘寒富'苹果叶片的叶肉和栅栏组织厚度、栅海比、气孔密度、基粒厚度、基粒片层数和比基粒片层数均显著高于‘东光'和‘富士';叶片的光合能力、表观量子效率、羧化效率和RuBP最大再生速率、叶绿素含量和比叶重较高;具有较高的初始荧光,较低的最大荧光产量、PSⅡ最大光化学效率和性能指数;‘寒富'苹果叶片光合作用的启动时间较短,大约在22min时就达到最大值(19.5μmol·m~(-2)·s~(-1));‘寒富'苹果具有较高的饱和光强、饱和CO_2和CO_2补偿点,较低的光补偿点,分别为(1665±20)μmol·m~(-2)·s~(-1)、(2961±41)μmol·mol~(-1)、(87.5±1.4)μmol·mol~(-1)、和(5.9±1.8)μmol·m~(-2)·s~(-1);年生长周期内,‘寒富'苹果叶片的可溶性糖和淀粉含量略低于‘东光',但明显高于‘富士'。
2不同发育状态的‘寒富'苹果叶片光合生理参数表现不同,功能叶和幼叶光合作用的启动时间较短,净光合速率大约在22min时就达到最大值(19.5μmol·m~(-2)·s~(-1)和12.29μmol·m~(-2)·s~(-1))。功能叶片的表观量子效率、气孔限制值和光合能力最大。功能叶的饱和光强、光补偿点和饱和CO_2均高于嫩叶和衰老叶,CO_2补偿点较低,分别为(2210±10.6)μmol·m~(-2)·s~(-1)、(28.5±2.8)μmol·m~(-2)·s~(-1)、(1222.7±42)μmol·mol~(-1)、和(85.6±1.5)μmol·mol~(-1),表明功能叶利用强光、高CO_2和低CO_2的能力较强。可溶性糖含量在叶片发育初期含量较高,随着开花和新梢旺长含量降低。新梢停长后可溶性糖和淀粉含量出现峰值,随着叶片的衰老下降。年生长周期内,‘寒富'苹果葡萄糖和山梨醇含量(44.587mg·ml~(-1)和32.287mg·ml~(-1))显著高于蔗糖和果糖含量(3.841mg·ml~(-1)和2.333mg·ml~(-1))。
3不同组织器官和枝类‘寒富'苹果的营养物质含量不同。叶片中的可溶性糖含量最高,韧皮部淀粉含量最高。秋梢叶片中可溶性糖含量、以及木质部的可溶性糖和淀粉含量高于春梢和封顶枝。养分再分配过程中,一年生枝韧皮部和木质部的淀粉含量、一年生枝木质部的可溶性糖含量高于其他级次枝,主干韧皮部的可溶性糖含量最高。
4‘寒富'苹果新梢不同节位叶片的光合生理参数表现不同。净光合速率呈由低至高的变化趋势,基部叶片的Pn较低,从基部第4节开始维持较高水平,并且各节位之间的差异较小。新梢中部11～16节位的叶片具有较低的初始荧光和最大荧光产量,PSⅡ最大光化学效率和性能指数均高于新梢基部和梢部叶片。
5覆草处理增加了‘寒富'节果的叶片厚度、栅栏组织厚度和栅海比,气孔密度、叶绿体数目、叶绿体长度、基粒片层数和比基粒片层数均呈现不同程度的增加,基粒类囊体排列紧密而整齐,叶绿体片层结构较发达;覆草处理增加了‘寒富'苹果叶片的光合色素含量,提高了‘寒富'苹果叶片的净光合速率、初始荧光、最大荧光产量和性能指数。覆草处理叶片、韧皮部和木质部的可溶性糖和淀粉含量明显高于清耕。
6弱光胁迫条件下,‘寒富'苹果叶片变薄,栅栏组织排列疏松,栅海比增加,叶绿体变小,基粒厚度明显增加,叶绿体基粒片层结构增厚,片层间隙增大,并出现大量的噬锇体;弱光胁迫条件下,‘寒富'苹果叶片的净光合速率、水分利用效率和气孔导度降低,叶片PSⅡ反应中心原初光能转化效率和PSⅡ的潜在活性提高;弱光降低了‘寒富'苹果叶片的光合能力、表观量子效率、羧化效率、气孔限制值和RuBP最大再生速率,饱和光强、光补偿点和饱和CO_2降低,CO_2补偿点升高,弱光下建造的‘寒富'苹果叶片突然转至强光下以后,光合作用的启动时间延长。弱光使可溶性糖含量增加,蔗糖磷酸合成酶活性上升,转化酶、山梨醇脱氢酶和山梨醇氧化酶活性下降,蔗糖合成酶活性趋于稳定,‘寒富'苹果具有喜光耐荫的特点。
7轻度干旱胁迫处理使‘寒富'苹果叶片的栅栏组织厚度增加,栅海比增大,上表皮出现一层失活的木栓组织,栅栏组织细胞中出现失活点。中度、重度干旱胁迫和淹水条件下,叶肉的栅栏组织和海绵组织细胞解体。适度水分胁迫增加了‘寒富'节果气孔导度和水分利用效率,对净光合速率没有影响,并且提高了PSⅡ反应中心原初光能转化效率和PSⅡ的潜在活性,随着胁迫程度的加重,‘寒富'苹果的光合参数和叶绿素荧光参数才受到不可逆转的影响;水分胁迫使‘寒富'苹果叶片内可溶性糖和淀粉含量降低,胁迫越严重,变化幅度越大。轻度干旱胁迫提高了‘寒富'苹果叶片中性转化酶、酸性转化酶和蔗糖合成酶活性,降低了蔗糖磷酸合成酶、山梨醇脱氢酶、山梨醇氧化酶活性。中度、重度干旱胁迫和淹水条件下,中性转化酶、酸性转化酶和蔗糖合成酶降低。‘寒富'苹果能忍耐轻度和中度干旱胁迫,抗旱性较强。
8高湿胁迫条件下,‘寒富'苹果叶片厚度增加,栅栏组织细胞明显增多,海绵组织结构被破坏,细胞解体,内部出现失活点。高湿处理降低了‘寒富'苹果净光合速率、PSⅡ反应中心原初光能转化效率和PSⅡ潜在活性,提高了初始荧光产量,但没有显著影响‘寒富'苹果叶片能量的分配;高湿胁迫使‘寒富'苹果叶片内可溶性糖含量升高,淀粉含量降低,提高了‘寒富'苹果叶片中性转化酶、酸性转化酶、蔗糖合成酶活性,降低了蔗糖磷酸合成酶、山梨醇脱氢酶、山梨醇氧化酶活性,‘寒富'苹果具有较强的耐湿性。
The breeding of 'Hanfu' apple played an important role for expanding the cultivation area and building the new superior industry belt of big fruit apple,but the researches of related physiological basis and the mechanism of technical measures are great shortage. Particularly,the researches on effects of cold weather conditions on related physiological basis of high-quality and big fruit apple has not been reported.In this paper,with 'Hanfu' apple and its parents 'Dongguang' and 'Fuji' apple as the materials,the photosynthetic physiology and carbon material metabolism of 'Hanfu' apple were evaluated preliminarily by the means of photosynthetic gas exchange and chlorophyll fluorescence analysis combining with tissue anatomy characteristic analysis and physiological and biochemical detection.The photosynthetic physiological and carbon material metabolism basis of 'Hanfu' apple under the conditions of a variety of stress,different soil management systems were studied for discussing the mechanism of photosynthetic physiological and carbon material metabolism, revealing the physiological basis of multiple resistance expression of 'Hanfu' apple in cold regions and providing a theoretical basis for high-quality cultivations and managements of 'Hanfu' apple in cold regions.The main results were as follows:
1 Compared with that of 'Dongguang' and 'Fuji'apple,the leaves of 'Hanfu' apple had higher thickness of mesophyll and the largest rate of palisade tissue and spongy tissue.The stomatal density,grana thickness,number of grana lamellae and specific lamella number of granum were significantly higher than its parents.The photosynthetic capacity,apparent quantum yield,carboxylation efficiency and maximum regeneration velocity of RuBP, chlorophyll content and specific leaf weight were higher than its parents.The leaves of 'Hanfu' apple had higher initial fluorescence,lower maximum fluorescence yield, PSⅡmaximum photochemical efficiency and performance index.The photosynthesis starting time of leaves in 'Hanfu' apple was shorter and the maximum(19.5μmol·m~(-2)·s~(-1)) was appeared at about 22 minutes.'Hanfu' apple had a higher saturation light intensity,Saturation CO_2,CO_2 compensation point and lower light compensation point,which respectively were (1665±20)μmol·m~(-2).s~(-1),(2961±41)μmol·mol~(-1),(87.5±1.4)μmol·mol~(-1),and (5.9±1.8)μmol·m~(-2)·s~(-1).During annual growth cycle,the eontents of soluble sugar and starch in 'Hanfu' apple were slightly lower than that of 'Dongguang' apple leaves,but significantly higher than that of 'Fuji' apple leaves.
2 The photosynthetic physiological parameters of different developmental leaves were different.The photosynthesis starting time of functional and young leaves were shorter start photosynthesis,the maximum(19.5μmol·m~(-2)·s~(-1) and 12.29μmol·m~(-2)·s~(-1)) were appeared at about 22 minutes.The apparent quantum efficiency and stomatal limitation value of functional leaves were significantly higher than that of young leaves and senescence leaf.The photosynthetic capacity of functional leaves and young leaves was no significant difference, but both were significantly higher than senescence leaf.The carboxylation efficiency and RuBP maximumn regeneration rate of leaves at different developmental stageshad no difference.The saturation light intensity,light compensation point and saturation CO_2 of functional leaves were higher than that of young leaves and senescence leaves whereas the CO_2 compensation points was lower,which respectively were(2210±10.6)μmol·m~(-2)·s~(-1), (28.5±2.8)μmol·m~(-2)·s~(-1),(1222.7±42)μmol·mol~(-1),and(85.6±1.5)μmol·mol~(-1).All showed that the functional leaves and the capacity of using high light,high and low CO_2.With the development of leaves,the net photosynthetic rate gradually increased,it reached the peak after leaf maturation and decreased rapidly with the aging of leaves.The soluble sugar contents of leaves were higher at the early stage of development,which decreased with the flowering and rapid growth of new shoots.After new shoots stopped growth,the soluble sugar and starch content reached peaks,which decreased with leaf senescence.During the annual growth cycle,the glucose and sorbitol content(44.587mg·ml~(-1) and 32.287mg·ml~(-1)) was significantly higher than that of sucrose and fructose content(3.841mg·ml~(-1) and 2.333mg·ml~(-1)) in 'Hanfu' apple.
3 The nutrient content of different types of tissues,organs and branches were different in 'Hanfu' apple.The soluble sugar content in leaves was the highest,and the starch content in phloem was the highest.The soluble sugar content in leaves,the soluble sugar and starch content in xylem of autumn shoots were higher than that of spring shoots and one-year-old top-cutting shoots.During the process of nutrient redistribution,the starch content in the one-year-old branch phloem and xylem,the soluble sugar in one-year-old branch xylem were the highest,and the soluble sugar content in trunk phloem was the highest.
4 The photosynthetic physiological parameters of leaves at different leaf node in 'Hanfu' apple were different.The net photosynthetic rate showed the trend of from low to high.The Pn of basal leaves was lower,it maintained a fairly high level from the base fourth node,and the difference among the nodes was smaller.The leaves at the 11th to 16th nodes had lower initial fluorescence and maximal fluorescence yield,the PSⅡmaximumn photochemical efficiency and performance index were higher than that of leaves at the base and tip of shoot.
5 Under planting and covering grass treatment,the thickness of leaves,palisade tissue and the rate of palisade tissue and spongy tissue,the stomatal density,the number of chloroplasts,chloroplast length,number of grana lamellae and specific lamella number of granum increased to some extent.The granum-thylakoid were arranged compactly and regularly with developed chloroplast lamellar structure.The photosynthetic pigment content, the net photosynthetic rate,the initial fluorescence,maximal fluorescence yield and the performance index improved.The soluble sugar and starch content of leaves,phloem and xylem increased.
6 Under low-light stress,the thickness of leaves and spongy tissue were thinned with loose palisade tissue,the rate of palisade tissue and spongy tissue and basal granule thickness increased.The size of chloroplast was significantly smaller,the number of grana lamellae and specific lamella number of granum decreased,the chloroplast grana lamellae structure was thickened,the lamellar gap was enlarged and a large number of plastoglobuli appeared. Under low light stress,the net photosynthetic rate,water use efficiency and stomatal conductance were reduced,and PSⅡconversion efficiency of primary light energy and the potential activity of PSⅡimproved.The photosynthetic capacity,apparent quantum yield, carboxylation efficiency,stomatal limitation value and the RuBP maximum regeneration rate, saturation light intensity,light compensation point and saturation CO_2 decreased with the weakness of light intensity,but the CO_2 compensation points increased.After the 'Hanfu' apple leaves constructed in low light transferred to the strong light suddenly,the photosynthesis starting time was prolonged.The soluble sugar content increased,the activity of sucrose phosphate synthase enhanced,but the activity of glutamine synthetase,invertase, sorbitol dehydrogenase and sorbitol oxidase declined,the activity of sucrose synthase maintained stability,which indicated that 'Hanfu' apple was a photophil and shade-tolerant plants.
7 Under light drought stress,the thickness of palisade tissue and the rate of palisade tissue and spongy tissue increased.A layer of inactivated phellem tissue appeared on the upper epidermis and the inactivation point appeared in the palisade tissue cells.In moderate and high drought stress and waterlogging conditions,the palisade tissue and spongy tissue cells in mesophyll disintegrated,a lot of broken cells and cavity appeared,and the proportion of xylem increased.Under moderate water stress,the stomatal conductance,water use efficiency increased and PSⅡconversion efficiency of primary light energy and the potential activity of PSⅡimproved,but the net photosynthetic rate had no effect.With the heavier of stress levels,the photosynthetic parameters and chlorophyll fluorescence parameters of 'Hanfu' apple had be affected irreversibly.In water stress conditions,the soluble sugar content increased,and the starch content reduced.The heavier of stress,the greater of change range.Under light drought stress,the neutral invertase,acid invertase and sucrose synthase activity improved,sucrose phosphate synthase,sorbitol dehydrogenase,sorbitol oxidase and glutamine synthetase activity reduced.In moderate and high drought stress and waterlogging conditions,neutral invertase,acid invertase and sucrose synthase activity reduced enhanced. 'hanfu' apple was a cultivar with strong drought resistance because it could tolerate light and moderate drought.
8 Under high humidity stress,the thickness of leaves and palisade tissue cells markedly increased.The structure of spongy tissue was damaged,mesophyll cells was disintegrated,a large number of cavity and inactivation point appeared.The net photosynthetic rate, PSⅡconversion efficiency of primary light energy and the potential activity of PSⅡreduced and the initial fluorescence yield increased,but the distribution of energy did not significantly be affected.The soluble sugar increased and starch contents decreased.The neutral invertase, acid invertase,sucrose synthase and glutamine synthetase activity improved,sucrose phosphate synthase,sorbitol dehydrogenase,sorbitol oxidase activity reduced.'hanfu' apple was a cultivar with strong high humidity resistance.
1‘寒富'苹果叶片的叶肉和栅栏组织厚度、栅海比、气孔密度、基粒厚度、基粒片层数和比基粒片层数均显著高于‘东光'和‘富士';叶片的光合能力、表观量子效率、羧化效率和RuBP最大再生速率、叶绿素含量和比叶重较高;具有较高的初始荧光,较低的最大荧光产量、PSⅡ最大光化学效率和性能指数;‘寒富'苹果叶片光合作用的启动时间较短,大约在22min时就达到最大值(19.5μmol·m~(-2)·s~(-1));‘寒富'苹果具有较高的饱和光强、饱和CO_2和CO_2补偿点,较低的光补偿点,分别为(1665±20)μmol·m~(-2)·s~(-1)、(2961±41)μmol·mol~(-1)、(87.5±1.4)μmol·mol~(-1)、和(5.9±1.8)μmol·m~(-2)·s~(-1);年生长周期内,‘寒富'苹果叶片的可溶性糖和淀粉含量略低于‘东光',但明显高于‘富士'。
2不同发育状态的‘寒富'苹果叶片光合生理参数表现不同,功能叶和幼叶光合作用的启动时间较短,净光合速率大约在22min时就达到最大值(19.5μmol·m~(-2)·s~(-1)和12.29μmol·m~(-2)·s~(-1))。功能叶片的表观量子效率、气孔限制值和光合能力最大。功能叶的饱和光强、光补偿点和饱和CO_2均高于嫩叶和衰老叶,CO_2补偿点较低,分别为(2210±10.6)μmol·m~(-2)·s~(-1)、(28.5±2.8)μmol·m~(-2)·s~(-1)、(1222.7±42)μmol·mol~(-1)、和(85.6±1.5)μmol·mol~(-1),表明功能叶利用强光、高CO_2和低CO_2的能力较强。可溶性糖含量在叶片发育初期含量较高,随着开花和新梢旺长含量降低。新梢停长后可溶性糖和淀粉含量出现峰值,随着叶片的衰老下降。年生长周期内,‘寒富'苹果葡萄糖和山梨醇含量(44.587mg·ml~(-1)和32.287mg·ml~(-1))显著高于蔗糖和果糖含量(3.841mg·ml~(-1)和2.333mg·ml~(-1))。
3不同组织器官和枝类‘寒富'苹果的营养物质含量不同。叶片中的可溶性糖含量最高,韧皮部淀粉含量最高。秋梢叶片中可溶性糖含量、以及木质部的可溶性糖和淀粉含量高于春梢和封顶枝。养分再分配过程中,一年生枝韧皮部和木质部的淀粉含量、一年生枝木质部的可溶性糖含量高于其他级次枝,主干韧皮部的可溶性糖含量最高。
4‘寒富'苹果新梢不同节位叶片的光合生理参数表现不同。净光合速率呈由低至高的变化趋势,基部叶片的Pn较低,从基部第4节开始维持较高水平,并且各节位之间的差异较小。新梢中部11～16节位的叶片具有较低的初始荧光和最大荧光产量,PSⅡ最大光化学效率和性能指数均高于新梢基部和梢部叶片。
5覆草处理增加了‘寒富'节果的叶片厚度、栅栏组织厚度和栅海比,气孔密度、叶绿体数目、叶绿体长度、基粒片层数和比基粒片层数均呈现不同程度的增加,基粒类囊体排列紧密而整齐,叶绿体片层结构较发达;覆草处理增加了‘寒富'苹果叶片的光合色素含量,提高了‘寒富'苹果叶片的净光合速率、初始荧光、最大荧光产量和性能指数。覆草处理叶片、韧皮部和木质部的可溶性糖和淀粉含量明显高于清耕。
6弱光胁迫条件下,‘寒富'苹果叶片变薄,栅栏组织排列疏松,栅海比增加,叶绿体变小,基粒厚度明显增加,叶绿体基粒片层结构增厚,片层间隙增大,并出现大量的噬锇体;弱光胁迫条件下,‘寒富'苹果叶片的净光合速率、水分利用效率和气孔导度降低,叶片PSⅡ反应中心原初光能转化效率和PSⅡ的潜在活性提高;弱光降低了‘寒富'苹果叶片的光合能力、表观量子效率、羧化效率、气孔限制值和RuBP最大再生速率,饱和光强、光补偿点和饱和CO_2降低,CO_2补偿点升高,弱光下建造的‘寒富'苹果叶片突然转至强光下以后,光合作用的启动时间延长。弱光使可溶性糖含量增加,蔗糖磷酸合成酶活性上升,转化酶、山梨醇脱氢酶和山梨醇氧化酶活性下降,蔗糖合成酶活性趋于稳定,‘寒富'苹果具有喜光耐荫的特点。
7轻度干旱胁迫处理使‘寒富'苹果叶片的栅栏组织厚度增加,栅海比增大,上表皮出现一层失活的木栓组织,栅栏组织细胞中出现失活点。中度、重度干旱胁迫和淹水条件下,叶肉的栅栏组织和海绵组织细胞解体。适度水分胁迫增加了‘寒富'节果气孔导度和水分利用效率,对净光合速率没有影响,并且提高了PSⅡ反应中心原初光能转化效率和PSⅡ的潜在活性,随着胁迫程度的加重,‘寒富'苹果的光合参数和叶绿素荧光参数才受到不可逆转的影响;水分胁迫使‘寒富'苹果叶片内可溶性糖和淀粉含量降低,胁迫越严重,变化幅度越大。轻度干旱胁迫提高了‘寒富'苹果叶片中性转化酶、酸性转化酶和蔗糖合成酶活性,降低了蔗糖磷酸合成酶、山梨醇脱氢酶、山梨醇氧化酶活性。中度、重度干旱胁迫和淹水条件下,中性转化酶、酸性转化酶和蔗糖合成酶降低。‘寒富'苹果能忍耐轻度和中度干旱胁迫,抗旱性较强。
8高湿胁迫条件下,‘寒富'苹果叶片厚度增加,栅栏组织细胞明显增多,海绵组织结构被破坏,细胞解体,内部出现失活点。高湿处理降低了‘寒富'苹果净光合速率、PSⅡ反应中心原初光能转化效率和PSⅡ潜在活性,提高了初始荧光产量,但没有显著影响‘寒富'苹果叶片能量的分配;高湿胁迫使‘寒富'苹果叶片内可溶性糖含量升高,淀粉含量降低,提高了‘寒富'苹果叶片中性转化酶、酸性转化酶、蔗糖合成酶活性,降低了蔗糖磷酸合成酶、山梨醇脱氢酶、山梨醇氧化酶活性,‘寒富'苹果具有较强的耐湿性。
The breeding of 'Hanfu' apple played an important role for expanding the cultivation area and building the new superior industry belt of big fruit apple,but the researches of related physiological basis and the mechanism of technical measures are great shortage. Particularly,the researches on effects of cold weather conditions on related physiological basis of high-quality and big fruit apple has not been reported.In this paper,with 'Hanfu' apple and its parents 'Dongguang' and 'Fuji' apple as the materials,the photosynthetic physiology and carbon material metabolism of 'Hanfu' apple were evaluated preliminarily by the means of photosynthetic gas exchange and chlorophyll fluorescence analysis combining with tissue anatomy characteristic analysis and physiological and biochemical detection.The photosynthetic physiological and carbon material metabolism basis of 'Hanfu' apple under the conditions of a variety of stress,different soil management systems were studied for discussing the mechanism of photosynthetic physiological and carbon material metabolism, revealing the physiological basis of multiple resistance expression of 'Hanfu' apple in cold regions and providing a theoretical basis for high-quality cultivations and managements of 'Hanfu' apple in cold regions.The main results were as follows:
1 Compared with that of 'Dongguang' and 'Fuji'apple,the leaves of 'Hanfu' apple had higher thickness of mesophyll and the largest rate of palisade tissue and spongy tissue.The stomatal density,grana thickness,number of grana lamellae and specific lamella number of granum were significantly higher than its parents.The photosynthetic capacity,apparent quantum yield,carboxylation efficiency and maximum regeneration velocity of RuBP, chlorophyll content and specific leaf weight were higher than its parents.The leaves of 'Hanfu' apple had higher initial fluorescence,lower maximum fluorescence yield, PSⅡmaximum photochemical efficiency and performance index.The photosynthesis starting time of leaves in 'Hanfu' apple was shorter and the maximum(19.5μmol·m~(-2)·s~(-1)) was appeared at about 22 minutes.'Hanfu' apple had a higher saturation light intensity,Saturation CO_2,CO_2 compensation point and lower light compensation point,which respectively were (1665±20)μmol·m~(-2).s~(-1),(2961±41)μmol·mol~(-1),(87.5±1.4)μmol·mol~(-1),and (5.9±1.8)μmol·m~(-2)·s~(-1).During annual growth cycle,the eontents of soluble sugar and starch in 'Hanfu' apple were slightly lower than that of 'Dongguang' apple leaves,but significantly higher than that of 'Fuji' apple leaves.
2 The photosynthetic physiological parameters of different developmental leaves were different.The photosynthesis starting time of functional and young leaves were shorter start photosynthesis,the maximum(19.5μmol·m~(-2)·s~(-1) and 12.29μmol·m~(-2)·s~(-1)) were appeared at about 22 minutes.The apparent quantum efficiency and stomatal limitation value of functional leaves were significantly higher than that of young leaves and senescence leaf.The photosynthetic capacity of functional leaves and young leaves was no significant difference, but both were significantly higher than senescence leaf.The carboxylation efficiency and RuBP maximumn regeneration rate of leaves at different developmental stageshad no difference.The saturation light intensity,light compensation point and saturation CO_2 of functional leaves were higher than that of young leaves and senescence leaves whereas the CO_2 compensation points was lower,which respectively were(2210±10.6)μmol·m~(-2)·s~(-1), (28.5±2.8)μmol·m~(-2)·s~(-1),(1222.7±42)μmol·mol~(-1),and(85.6±1.5)μmol·mol~(-1).All showed that the functional leaves and the capacity of using high light,high and low CO_2.With the development of leaves,the net photosynthetic rate gradually increased,it reached the peak after leaf maturation and decreased rapidly with the aging of leaves.The soluble sugar contents of leaves were higher at the early stage of development,which decreased with the flowering and rapid growth of new shoots.After new shoots stopped growth,the soluble sugar and starch content reached peaks,which decreased with leaf senescence.During the annual growth cycle,the glucose and sorbitol content(44.587mg·ml~(-1) and 32.287mg·ml~(-1)) was significantly higher than that of sucrose and fructose content(3.841mg·ml~(-1) and 2.333mg·ml~(-1)) in 'Hanfu' apple.
3 The nutrient content of different types of tissues,organs and branches were different in 'Hanfu' apple.The soluble sugar content in leaves was the highest,and the starch content in phloem was the highest.The soluble sugar content in leaves,the soluble sugar and starch content in xylem of autumn shoots were higher than that of spring shoots and one-year-old top-cutting shoots.During the process of nutrient redistribution,the starch content in the one-year-old branch phloem and xylem,the soluble sugar in one-year-old branch xylem were the highest,and the soluble sugar content in trunk phloem was the highest.
4 The photosynthetic physiological parameters of leaves at different leaf node in 'Hanfu' apple were different.The net photosynthetic rate showed the trend of from low to high.The Pn of basal leaves was lower,it maintained a fairly high level from the base fourth node,and the difference among the nodes was smaller.The leaves at the 11th to 16th nodes had lower initial fluorescence and maximal fluorescence yield,the PSⅡmaximumn photochemical efficiency and performance index were higher than that of leaves at the base and tip of shoot.
5 Under planting and covering grass treatment,the thickness of leaves,palisade tissue and the rate of palisade tissue and spongy tissue,the stomatal density,the number of chloroplasts,chloroplast length,number of grana lamellae and specific lamella number of granum increased to some extent.The granum-thylakoid were arranged compactly and regularly with developed chloroplast lamellar structure.The photosynthetic pigment content, the net photosynthetic rate,the initial fluorescence,maximal fluorescence yield and the performance index improved.The soluble sugar and starch content of leaves,phloem and xylem increased.
6 Under low-light stress,the thickness of leaves and spongy tissue were thinned with loose palisade tissue,the rate of palisade tissue and spongy tissue and basal granule thickness increased.The size of chloroplast was significantly smaller,the number of grana lamellae and specific lamella number of granum decreased,the chloroplast grana lamellae structure was thickened,the lamellar gap was enlarged and a large number of plastoglobuli appeared. Under low light stress,the net photosynthetic rate,water use efficiency and stomatal conductance were reduced,and PSⅡconversion efficiency of primary light energy and the potential activity of PSⅡimproved.The photosynthetic capacity,apparent quantum yield, carboxylation efficiency,stomatal limitation value and the RuBP maximum regeneration rate, saturation light intensity,light compensation point and saturation CO_2 decreased with the weakness of light intensity,but the CO_2 compensation points increased.After the 'Hanfu' apple leaves constructed in low light transferred to the strong light suddenly,the photosynthesis starting time was prolonged.The soluble sugar content increased,the activity of sucrose phosphate synthase enhanced,but the activity of glutamine synthetase,invertase, sorbitol dehydrogenase and sorbitol oxidase declined,the activity of sucrose synthase maintained stability,which indicated that 'Hanfu' apple was a photophil and shade-tolerant plants.
7 Under light drought stress,the thickness of palisade tissue and the rate of palisade tissue and spongy tissue increased.A layer of inactivated phellem tissue appeared on the upper epidermis and the inactivation point appeared in the palisade tissue cells.In moderate and high drought stress and waterlogging conditions,the palisade tissue and spongy tissue cells in mesophyll disintegrated,a lot of broken cells and cavity appeared,and the proportion of xylem increased.Under moderate water stress,the stomatal conductance,water use efficiency increased and PSⅡconversion efficiency of primary light energy and the potential activity of PSⅡimproved,but the net photosynthetic rate had no effect.With the heavier of stress levels,the photosynthetic parameters and chlorophyll fluorescence parameters of 'Hanfu' apple had be affected irreversibly.In water stress conditions,the soluble sugar content increased,and the starch content reduced.The heavier of stress,the greater of change range.Under light drought stress,the neutral invertase,acid invertase and sucrose synthase activity improved,sucrose phosphate synthase,sorbitol dehydrogenase,sorbitol oxidase and glutamine synthetase activity reduced.In moderate and high drought stress and waterlogging conditions,neutral invertase,acid invertase and sucrose synthase activity reduced enhanced. 'hanfu' apple was a cultivar with strong drought resistance because it could tolerate light and moderate drought.
8 Under high humidity stress,the thickness of leaves and palisade tissue cells markedly increased.The structure of spongy tissue was damaged,mesophyll cells was disintegrated,a large number of cavity and inactivation point appeared.The net photosynthetic rate, PSⅡconversion efficiency of primary light energy and the potential activity of PSⅡreduced and the initial fluorescence yield increased,but the distribution of energy did not significantly be affected.The soluble sugar increased and starch contents decreased.The neutral invertase, acid invertase,sucrose synthase and glutamine synthetase activity improved,sucrose phosphate synthase,sorbitol dehydrogenase,sorbitol oxidase activity reduced.'hanfu' apple was a cultivar with strong high humidity resistance.
引文
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