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沙棘(Hippophae rhamnoides L.)属胡颓子科沙棘属植物,生态适应性强,果实营养成分丰富[1],是重要的经济生态两用树种。我国是世界上沙棘资源最丰富的国家之一,其中,中国沙棘亚种(Hippophae rhamnoides subsp. L. sinensis Rousi)是我国沙棘资源中分布面积最广的类型[2-3]。对中国沙棘生理和生长结实的研究有利于深入了解中国沙棘的生长结实习性与生理过程的关系。
树冠不同部位的光照条件、温度、湿度、通风状况等微气候条件存在较大差异[4-5],通常情况下上部枝条光照情况良好,枝条直立;随枝条着生位置的下移,枝条光照条件变差,且开张角度变大。前人研究表明:树冠不同部位枝条的生长、水碳代谢、结实性状等存在较大差异[6-7],但前人的研究多从树形、遗传特性及光合作用的角度对其结实特性的影响进行了分析[8-10],且多针对苹果、梨等大宗水果进行,而笔者的前期观察表明,沙棘的生长结实性状与苹果、梨等树种存在较大差异,且目前对沙棘树冠不同位置枝条的生长结实性状及其影响机制尚不了解。
水碳代谢是树木生长和结实的基础,前人研究表明,树木的水分状况、碳素营养的摄取和分配会影响到植物的生长以及碳平衡[11-12],也会影响到树木的花芽分化、开花结实和果实品质[13]。另外,树木的冠层特征以及同一棵树的不同部位,受其几何结构和枝叶分布的影响,其光通量、光能截获、光合作用以及蒸腾作用等也会产生差异[14],从而对树木的生长发育产生影响;但目前对沙棘枝条位置造成的水碳代谢差异、不同冠层结构和枝叶生长对其生长和结实性状的影响尚不了解。
本研究以中国沙棘为研究对象,通过测定树冠上、中、下部枝条的水分状况、光合作用、不同部位的非结构性碳水化合物含量、生长性状、结实性状和果实营养成分,比较分析沙棘树冠不同部位枝条的水碳代谢和生长结实性状的差别,并探讨树冠不同位置枝条的水碳代谢与枝条生长、结实性状和果实营养成分的关系,研究着生部位对沙棘枝条水碳代谢和生长结实的影响,研究结果有助于深入了解树冠位置差异对沙棘碳素分配和生长结实的影响,在生产上为沙棘栽培技术的制定提供理论依据。
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图1表明:沙棘树冠上、中、下部枝条的凌晨水势差异不显著,下部枝条的正午水势显著高于中部和上部枝条(p<0.05)。中部枝条的凌晨PLC显著低于上部和下部枝条的(p<0.05),下部枝条的正午PLC显著高于上部和中部枝条的(p<0.05),上部和中部枝条的正午PLC间差异不显著。
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图2表明:树冠上、中、下部枝条的茎面积比导水率间均差异显著(p<0.05),从大到小的顺序为:上部>中部>下部;中部枝条和下部枝条的茎面积比导水率比上部枝条分别降低了30.8%、55.5%。树冠上部枝条的叶面积比导水率显著高于中部枝条和下部枝条(p<0.05),但中部枝条和下部枝条的叶面积比导水率差异不显著,中部枝条和下部枝条的叶面积比导水率比上部枝条分别下降了44.8%和45.8%。
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图3表明:树冠上、中、下部叶片的净光合速率、水分利用效率间均差异显著(p<0.05),从大到小的顺序为上部>中部>下部,中部和下部枝条的光合速率比上部枝条的分别下降了22.1%和55.8%,上部枝条和中部枝条叶片的气孔导度和蒸腾速率均差异不显著,但均显著大于下部叶片(p<0.05)。
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图4表明:中部枝条木质部可溶性糖含量显著高于上部枝条(p<0.05),也略高于下部枝条,不同部位枝条韧皮部可溶性糖含量差异不显著,上部和中部叶片可溶性糖含量均显著大于下部叶片(p<0.05),但上部和中部叶片可溶性糖含量差异不显著。中部枝条木质部淀粉含量显著大于上部和下部(p<0.05),枝条韧皮部淀粉含量间差异不显著,上部和中部叶片的淀粉含量显著大于下部叶片(p<0.05)。中部枝条木质部的非结构性碳水化合物(NSC)含量显著大于上部枝条,略大于下部枝条,但差异不显著(p=0.07),3个部位枝条韧皮部间的NSC含量差异不显著,上部和中部叶片的NSC含量差异不显著,但均显著大于下部叶片(p<0.05)。
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表1表明:上部枝条的10 cm芽数、纯花芽比例显著大于下部,叶芽比例显著小于下部(p<0.05),混合芽比例在不同部位间差异不显著,上、中、下部枝条叶芽比例间均差异显著(p<0.05),从大到小的顺序为下部>中部>上部,上部和中部枝条的花芽纵径、花芽横径、发芽率显著大于下部枝条(p<0.05)。
表 1 着生位置对枝条、芽、叶片形态的影响
Table 1. The influence of the position of implantation on the morphology of branches,buds and leaves
测量指标
Measurement index上部枝条
Top branch中部枝条
Middle branch下部枝条
Bottom branch芽
Bud10 cm芽数 Buds number of 10 cm branches/个 12.6±1.4 a 12.1±0.9 ab 10.3±1.9 b 纯花芽比例 Flower bud ratio/% 74±8 a 67±8 a 40±2 b 混合芽比例 Mixed bud ratio/% 15±6 a 12±5 a 17±2 a 叶芽比例 Leaf bud ratio/% 11±2 c 21±5 b 43±2 a 花芽纵径 Flower bud longitudinal diameter/mm 2.45±0.08 a 2.37±0.16 a 1.93±0.12 b 花芽横径 Flower bud transverse diameter/mm 3.35±0.18 a 3.53±0.0.11 a 2.86±0.18 b 发芽率 germination percentage/% 91.2±5.3 a 92.2±6.0 a 77.2±8.2 b 枝
Branch2年生枝条长度 Biennial branch length/cm 66.0±3.4 a 57.8±3.9 b 46±7.4 c 2年生枝条基径 Biennial branch base diameter/mm 9.8±1.1 a 8.2±0.9 b 7.1±1.6 c 新梢数量 Number of new shoots /个 10.5±2.6 a 11.5±2.9 a 5.2±2.6 b >10 cm新梢数量 Number of new shoots greater than 10 cm/个 4.4±1.9 a 4.4±2.3 a 5.6±2.7 a >10 cm新梢基径 Shoot base diameter greater than 10 cm/mm 3.7±0.5 a 3.7±0.5 a 2.8±0.4 b >10 cm新梢长度 Shoot length greater than 10 cm/cm 30.0±6.8 a 33.7±5.5 a 20.3±2.2 b 叶
Leaf叶面积 Leaf area/mm2 378.2±31.0 a 386.4±19.8 a 309.8±23.2 b 比叶质量 Specific leaf weight/(g·m−2) 114.8±12.3 a 117.7±15.3 a 101.5±22.4 b 2年生枝条的长度和基径从大到小的顺序为上部>中部>下部,且均差异显著(p<0.05)。上部和中部单个2年生枝条上生长的新梢总数显著大于下部枝条(p<0.05),但长度大于10 cm新梢数量间差异不显著。上部和中部枝条的>10 cm新梢长度和新梢基径均显著大于下部枝条(p<0.05)。
上部和中部枝条的叶面积和比叶质量均显著大于下部(p<0.05),但上部和中部枝条的叶面积和比叶质量间差异不显著,下部枝条叶片的叶面积为上部叶片的81.9%,比叶质量为上部叶片的88.4%。
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表2表明:中部枝条的10 cm结果数、百果质量、果实横径、果实纵径、果柄长度均显著大于下部枝条(p<0.05),上部枝条的10 cm结果数和百果质量也显著大于下部枝条(p<0.05),但上部枝条的果实纵径、果实横径及果柄长度与下部枝条无显著差异,且上部枝条的百果质量、果实纵径、果实横径显著性低于中部枝条(p<0.05)。上部枝条的坐果率显著大于下部枝条(p<0.05),也略高于中部枝条,但差异不显著。上部和中部枝条果实的可溶性固形物含量显著高于下部果实(p<0.05);上、中、下部枝条果实的总黄酮、维生素C和可滴定酸含量均差异不显著。
表 2 着生位置对枝条结实性状和果实营养成分的影响
Table 2. The influence of the position of implantation on branch fruiting characters and fruit nutritional components
测量指标
Measurement index上部枝条
Top branch中部枝条
Middle branch下部枝条
Bottom branch结实性状
Seed setting character10 cm结果数 Fruits number of 10 cm branches/个 23.2±3.6 a 26.2±1.6 a 12.3±3.7 b 百果质量 Weight per 100 fruit/g 11.6±0.3 b 12.4±0.2 a 10.9±0.2 c 果实纵径 Fruit longitudinal diameter/mm 5.36±0.30 b 5.70±0.27 a 5.19±0.28 b 果实横径 Fruit transverse diameter/mm 6.56±0.30 b 6.90±0.29 a 6.49±0.26 b 果柄长度 Fruit stalk length/mm 1.96±0.09 ab 2.04±0.17 a 1.89±0.18 b 坐果率 Fruit setting rate/% 73.7±7.2 a 64.3±5.6 ab 52.9±2.0 b 果实营养成分
Fruit nutrients可溶性固形物 Soluble solids/% 13.97±0.96 a 14.15±0.08 a 9.38±0.15 b 总黄酮 Flavonoids/(mg·g−1) 11.1±0.7 a 10.7±0.2 a 10.7±1.7 a 维生素C Vitamin C/(mg·g−1) 5.87±0.54 a 6.20±0.50 a 6.00±0.60 a 可滴定酸 Titratable acid/% 2.18±0.12 a 2.39±0.25 a 2.35±0.14 a
中国沙棘树冠上中下部枝条的水碳代谢与生长结实性状研究
Correlation between Water-carbon Metabolism and Growth and Fruiting Traits of Upper, Middle and Lower Branches in the Crown of Hippophae rhamnoides subsp. sinensis Rousi
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摘要:
目的 为了解沙棘树冠不同位置枝条的水碳代谢与生长结实的关系,也为制定针对沙棘树形培养的栽培技术措施提供一定的理论依据。 方法 本研究选取中国沙棘树冠上部、中部、下部的2年生枝条为研究对象,比较其水分状况、光合作用、非结构性碳水化合物(NSC)含量、枝叶生长、结实性状和果实营养成分。 结果 树冠不同部位枝条凌晨水势差异不显著,下部枝条正午水势显著高于上部和中部枝条;中部枝条凌晨导水损失率(PLC)显著低于上部和下部枝条,下部枝条正午PLC显著高于上部和中部枝条;上部枝条的茎、叶面积比导水率显著高于中部和下部枝条;净光合速率按从大到小的顺序为上部枝条>中部枝条>下部枝条;中部枝条枝条木质部NSC含量显著高于上部枝条,但二者与下部枝条差异都不显著,各位置间枝条韧皮部NSC含量无显著性差异,上部和中部枝条的叶片NSC含量显著性高于下部枝条;下部枝条的花芽纵横径、纯花芽比例、发芽率、新梢数量、新梢基径、叶面积、比叶质量均显著低于上部和中部枝条,下部枝条的果实纵横径、坐果率、可溶性固形物含量也低于上部和中部枝条,上部枝条的枝长、基径显著高于中部枝条,中部枝条的百果质量和果实纵横径显著高于上部枝条,不同部位枝条总黄酮、维生素C和可滴定酸含量差异不显著。 结论 下部枝条受遮光的影响较大,明显限制光合产物摄取和生长,并较大程度的影响了其结实性状,上部枝条在生长上占有优势,中部枝条在结实性状上占有优势。因此,中国沙棘的树形培养应充分考虑其光照条件,及时清除下部低光照区域枝条;同时注意营养生长和生殖生长的平衡,充分利用树冠上部和中部枝条结实是提高沙棘产量和品质的重要措施。 Abstract:Objective To understand the relationship between water and carbon metabolism and growth and fruiting of branches at different positions in the crown of Hippophae rhamnoides for providing a certain theoretical basis for the formulation of cultivation technical measures for Hippophae rhamnoides tree shape culture. Method According to the two-year-old branches in the upper, middle and lower crown of Hippophae rhamnoides, water status, photosynthesis, non-structural carbohydrate (NSC) content, branch and leaf growth, fruiting traits and fruit nutrients were compared. Result The results showed that there was no significant difference in the pre-dawn water potential of branches in different parts of the crown, and the water potential of the bottom branches at midday was significantly higher than that of the upper and middle branches. The percentage loss of conductivity (PLC) of the middle branch in pre-dawn was significantly lower than that of upper and lower branches, and the PLC of the bottom branch at midday was significantly higher than that of top and middle branches. The stem and leaf area specific hydraulic conductivity of upper branches was significantly higher than that of middle and lower branches. The net photosynthetic rate in the top branch was the largest, followed by middle branch and bottom branch. NSC content in xylem of middle branches was significantly higher than that of upper branches, but there was no significant difference between the upper and lower branches, as well as between the middle and lower branches. There was no significant difference in NSC content in phloem among branches, and the NSC content in leaves of upper and middle branches was significantly higher than that of lower branches. The longitudinal and transverse diameter of flower buds, the proportion of pure flower buds, the germination rate, the number of new shoots, the basal diameter of new shoots, the leaf area and the specific leaf mass of the bottom branches were significantly lower than those of the upper and middle branches. The longitudinal and transverse diameter of fruit, fruit setting rate and soluble solid content of the bottom branches were both lower than those of the upper and middle branches. The branch length and base diameter of the upper branches are significantly higher than that of the middle branch. The fruit quality and fruit vertical and horizontal diameter of the middle branch were significantly higher than those of the upper branch. There was no significant difference in the contents of flavonoids, vitamin C and titratable acid among different branches. Conclusion The lower branches are greatly affected by shading, which significantly restricts the uptake and growth of photosynthetic products, and affects their fruiting traits. The upper branches are dominant in growth, and the middle branches are dominant in fruiting traits. Therefore, the tree-shaped cultivation of Hippophae rhamnoides should take full account of its light conditions, and timely remove the branches in the lower light area. At the same time, we should pay attention to the balance between nutritional growth and reproductive growth, and making full use of the upper and middle branches of the crownfor improving the yield and quality of Hippophae rhamnoides. -
表 1 着生位置对枝条、芽、叶片形态的影响
Table 1. The influence of the position of implantation on the morphology of branches,buds and leaves
测量指标
Measurement index上部枝条
Top branch中部枝条
Middle branch下部枝条
Bottom branch芽
Bud10 cm芽数 Buds number of 10 cm branches/个 12.6±1.4 a 12.1±0.9 ab 10.3±1.9 b 纯花芽比例 Flower bud ratio/% 74±8 a 67±8 a 40±2 b 混合芽比例 Mixed bud ratio/% 15±6 a 12±5 a 17±2 a 叶芽比例 Leaf bud ratio/% 11±2 c 21±5 b 43±2 a 花芽纵径 Flower bud longitudinal diameter/mm 2.45±0.08 a 2.37±0.16 a 1.93±0.12 b 花芽横径 Flower bud transverse diameter/mm 3.35±0.18 a 3.53±0.0.11 a 2.86±0.18 b 发芽率 germination percentage/% 91.2±5.3 a 92.2±6.0 a 77.2±8.2 b 枝
Branch2年生枝条长度 Biennial branch length/cm 66.0±3.4 a 57.8±3.9 b 46±7.4 c 2年生枝条基径 Biennial branch base diameter/mm 9.8±1.1 a 8.2±0.9 b 7.1±1.6 c 新梢数量 Number of new shoots /个 10.5±2.6 a 11.5±2.9 a 5.2±2.6 b >10 cm新梢数量 Number of new shoots greater than 10 cm/个 4.4±1.9 a 4.4±2.3 a 5.6±2.7 a >10 cm新梢基径 Shoot base diameter greater than 10 cm/mm 3.7±0.5 a 3.7±0.5 a 2.8±0.4 b >10 cm新梢长度 Shoot length greater than 10 cm/cm 30.0±6.8 a 33.7±5.5 a 20.3±2.2 b 叶
Leaf叶面积 Leaf area/mm2 378.2±31.0 a 386.4±19.8 a 309.8±23.2 b 比叶质量 Specific leaf weight/(g·m−2) 114.8±12.3 a 117.7±15.3 a 101.5±22.4 b 表 2 着生位置对枝条结实性状和果实营养成分的影响
Table 2. The influence of the position of implantation on branch fruiting characters and fruit nutritional components
测量指标
Measurement index上部枝条
Top branch中部枝条
Middle branch下部枝条
Bottom branch结实性状
Seed setting character10 cm结果数 Fruits number of 10 cm branches/个 23.2±3.6 a 26.2±1.6 a 12.3±3.7 b 百果质量 Weight per 100 fruit/g 11.6±0.3 b 12.4±0.2 a 10.9±0.2 c 果实纵径 Fruit longitudinal diameter/mm 5.36±0.30 b 5.70±0.27 a 5.19±0.28 b 果实横径 Fruit transverse diameter/mm 6.56±0.30 b 6.90±0.29 a 6.49±0.26 b 果柄长度 Fruit stalk length/mm 1.96±0.09 ab 2.04±0.17 a 1.89±0.18 b 坐果率 Fruit setting rate/% 73.7±7.2 a 64.3±5.6 ab 52.9±2.0 b 果实营养成分
Fruit nutrients可溶性固形物 Soluble solids/% 13.97±0.96 a 14.15±0.08 a 9.38±0.15 b 总黄酮 Flavonoids/(mg·g−1) 11.1±0.7 a 10.7±0.2 a 10.7±1.7 a 维生素C Vitamin C/(mg·g−1) 5.87±0.54 a 6.20±0.50 a 6.00±0.60 a 可滴定酸 Titratable acid/% 2.18±0.12 a 2.39±0.25 a 2.35±0.14 a -
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