毛竹出笋后快速生长期内茎秆中光合色素和光合酶活性的变化

doi:10.3724/SP.J.1258.2012.00456

摘要/Abstract

摘要：

为了探讨毛竹(Phyllostachys pubescens)非同化器官茎秆的光合特性, 测定了毛竹出笋后快速生长期内(2011年4月13日到6月2日)的光合色素含量以及光合酶活性, 并通过激光共聚焦显微镜对其叶绿体分布进行了观察。结果表明: 毛竹茎秆中的叶绿体主要集中分布在表皮以下的基本组织中, 此外维管束鞘周围的细胞内也存在大量的叶绿体, 此特征类似于C₄植物的花环结构。在毛竹出笋后快速生长期内, 随着茎秆不断生长, 叶绿素a、叶绿素b和类胡萝卜素含量均极显著(p < 0.01)增加。在出笋10天时, 茎秆中核酮糖-1,5-二磷酸羧化酶/加氧酶(Rubisco)活性、磷酸烯醇式丙酮酸羧化酶(PEPC)活性和NADP-苹果酸酶(NADP-ME)活性最高, 之后随茎秆生长逐渐降低, 生长30天时酶活性与10天时相比分别降低了88.55% (p < 0.01)、77.46% (p < 0.01)和72.50% (p < 0.01), 而PEPC/Rubisco比值则随茎秆生长逐渐增加, 30天时比值达到12.83, 明显高于C₃植物。这表明毛竹茎秆内可能存在C₄光合途径, 此途径有利于毛竹提高光合效率, 进而促进其出笋后的快速生长。

关键词: 叶绿体, 光合酶, 光合色素, 毛竹, 茎秆

Abstract:

Aims Our objective was to reveal photosynthetic characters in Phyllostachys pubescens stems. We determined the distribution of chloroplasts and detected changes of pigment content and photosynthetic enzyme activity in the stems during their rapid growth stage.
Methods We detected the pigment content and photosynthetic enzyme activity by the colorimetric method and observed the distribution of chloroplasts using a laser scanning confocal microscope.
Important findings Chloroplasts were mainly distributed in the ground tissues under the epidermis and in cells around the vascular bundle, similar to Kranz anatomy in C₄ plants. The content of chlorophyll a, chlorophyll b and carotenoid was significantly (p < 0.01) increased with the growth of the stems of P. pubescens. The activities of ribulose-1, 5-bisphosphate carboxylase change/add oxygen enzymes (Rubisco), phosphoenolpyruvate carboxylase (PEPC) and nicotinamide-adenine dinucleotide phosphate-malic enzyme (NADP-ME) in the stems reached the highest level after 10 days, and they declined gradually with the growth of the stems. After 30 days, the activities decreased by 88.55% (p < 0.01), 77.46% (p < 0.01) and 72.50% (p < 0.01), respectively, compared with after 10 days. The ratio of PEPC/Rubisco increased gradually and reached 12.83 after 30 days, which was markedly higher than that in C₃ plants. These results indicated that there was C₄ photosynthetic pathway in the stems, and the pathway may play an important role in the efficient photosynthesis and rapid growth of P. pubescens.

Key words: chloroplast, photosynthetic enzyme, photosynthetic pigment, Phyllostachys pubescens, stem

王星星, 刘琳, 张洁, 王玉魁, 温国胜, 高荣孚, 高岩, 张汝民. 毛竹出笋后快速生长期内茎秆中光合色素和光合酶活性的变化. 植物生态学报, 2012, 36(5): 456-462. DOI: 10.3724/SP.J.1258.2012.00456

WANG Xing-Xing, LIU Lin, ZHANG Jie, WANG Yu-Kui, WEN Guo-Sheng, GAO Rong-Fu, GAO Yan, ZHANG Ru-Min. Changes of photosynthetic pigment and photosynthetic enzyme activity in stems of Phyllostachys pubescens during rapid growth stage after shooting. Chinese Journal of Plant Ecology, 2012, 36(5): 456-462. DOI: 10.3724/SP.J.1258.2012.00456

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链接本文: https://www.plant-ecology.com/CN/10.3724/SP.J.1258.2012.00456

https://www.plant-ecology.com/CN/Y2012/V36/I5/456

图/表 5

图1 毛竹茎秆中的叶绿体分布。A, 表皮下的基本组织。B, 维管束周围的基本组织。Chl, 叶绿体(红色); E, 表皮; F, 纤维; Gr, 基本组织; Va, 维管束。

Fig. 1 Distribution of chloroplast in the stems of Phyllostachys pubescens. A, Ground tissue under epidermis. B, Ground tissue around the vascular bundle. Chl, chloroplast (red); E, epidermis; F, fiber; Gr, ground tissue; Va, vascular bundle.

表1 毛竹出笋后快速生长期茎秆中色素含量的变化(平均值±标准偏差)

Table 1 Changes of pigment contents in the stems of Phyllostachys pubescens under rapid growth stage after shooting (μg·g-1 FW) (mean ± SD)

出笋后的生长天数 Growth days after shooting (d)	叶绿素a Chlorophyll a	叶绿素b Chlorophyll b	叶绿素总量 Total chlorophyll	类胡萝卜素Carotenoids	叶绿素a/b Chlorophyll a/b
10	3.9 ± 0.6^eE	3.3 ± 1.2^eE	7.2 ± 1.8^eE	1.0 ± 0.3^eE	1.2 ± 0.8 ^bB
20	5.5 ± 0.8^eE	4.2 ± 1.2^eDE	9.7 ± 2.1^eE	0.9 ± 0.3^eE	1.3 ± 0.4^bB
30	17.7 ± 1.6^dD	7.2 ± 0.7^dD	24.9 ± 0.9^dD	6.5 ± 0.6^dD	2.5 ± 0.5^aA
40	135.3 ± 0.1^cC	65.3 ± 2.2^cC	200.7 ± 2.2^cC	42.1 ± 0.1^cC	2.0 ± 0.2^aA
50	179.8 ± 0.9^bB	94.6 ± 1.7^bB	274.4 ± 1.9^bB	46.2 ± 0.3^bB	2.0 ± 0.3^aA
60	269.6 ± 4.0^aA	128.0 ± 1.2^aA	397.6 ± 5.2^aA	49.8 ± 3.6^aA	2.1 ± 0.1^aA

图2 毛竹茎秆中核酮糖-1,5-二磷酸羧化酶/加氧酶(Ru- bisco)活性的变化(平均值±标准偏差)。

Fig. 2 Changes of ribulose-1,5-bisphosphate carboxylase change/add oxygen enzymes (Rubisco) activity in the stems of Phyllostachys pubescens (mean ± SD).

图3 毛竹茎秆中磷酸烯醇式丙酮酸羧化酶(PEPC)和烟酰胺腺嘌呤二核苷酸磷酸-苹果酸酶(NADP-ME)活性的变化(平均值±标准偏差)。

Fig. 3 Changes of phosphoenolpyruvate carboxylase (PEPC) and nicotinamide-adenine dinucleotide phosphate-malic enzyme (NADP-ME) activities in the stems of Phyllostachys pubescens (mean ± SD).

图4 毛竹茎秆中磷酸烯醇式丙酮酸羧化酶和核酮糖-1,5-二磷酸羧化酶/加氧酶比值(PEPC/Rubisco) (平均值±标准偏差)。

Fig. 4 Ratio of phosphoenolpyruvate carboxylase and ribulose-1,5-bisphosphate carboxylase change/add oxygen enzymes (PEPC/Rubisco) in the stems of Phyllostachys pubescens (mean ± SD).

参考文献 31

[1]	Aschan G, Pfanz H, Vodnik D, Batič F (2005). Photosynthetic performance of vegetative and reproductive structures of green hellebore (Helleborus viridis L. agg.). Photosynthetica, 43, 55-64.
[2]	Aschan G, Wittman C, Pfanz H (2001). Age-dependent bark photosynthesis of aspen twigs. Trees, 15, 431-437.
[3]	Berveiller D, Damesin C (2008). Carbon assimilation by tree stems: potential involvement of phosphoenolpyruvate carboxylase. Trees, 22, 149-157.
[4]	Berveiller D, Kierzkowski D, Damesin C (2007). Interspecific variability of stem photosynthesis among tree species. Tree Physiology, 27, 53-61.
[5]	Cemusak LA, Hutley LB (2011). Stable isotopes reveal the contribution of corticular photosynthesis to growth in branches of Eucalyptus miniata. Plant Physiology, 155, 515-523. URL PMID
[6]	Chen JH (陈建华), Mao D (毛丹), Ma ZY (马宗艳), Gong JH (龚建海) (2006). Physiological characteristics of leaves of bamboo Phyllostachys pubescens. Journal of Central South Forestry University (中南林学院学报), 26(6), 76-80. (in Chinese with English abstract)
[7]	Damesin C (2003). Respiration and photosynthesis characteristics of current-year stems of Fagus sylvatica: from the seasonal pattern to an annual balance. New Phytologist, 158, 465-475.
[8]	Häusler RE, Rademacher T, Li J, Lipka V, Fischer KL, Schubert S, Kreuzaler F, Hirsch HJ (2001). Single and double overexpression of C₄-cycle genes had differential effects on the pattern of endogenous enzymes, attenuation of photorespiration and on contents of UV protectants in transgenic potato and tobacco plants. Journal of Experimental Botany, 52, 1785-1803.
[9]	Hibberd JM, Quick WP (2002). Characteristics of C₄ photosynthesis in stems and petioles of flowering plants. Nature, 415, 451-454. DOI URL PMID
[10]	Holl W (1974). Dark CO₂ fixation by cell-free preparations of the wood of Robinia pseudoacacia. Canadian Journal of Botany, 52, 727-734.
[11]	Ivanov AG, Krol M, Sveshnikov D, Malmberg G, Gardeström P, Hurry V, Öquist Q, Huner NPA (2006). Characteriza- tion of the photosynthetic apparatus in cortical bark chlorenchyma of Scots pine. Planta, 223, 1165-1177.
[12]	Jiang ZS (姜振升), Sun XQ (孙晓琦), Ai XZ (艾希珍), Wang ML (王美玲), Bi HG (毕焕改), Wang HT (王洪涛) (2010). Responses of Rubisco and Rubisco activase in cucumber seedlings to low temperature and weak light. Chinese Journal of Applied Ecology (应用生态学报), 21, 2045-2050. (in Chinese with English abstract)
[13]	Kauppi A (1991). Seasonal fluctuations in chlorophyll content in birch stems with special reference to bark thickness and light transmission, a comparison between sprouts and seedlings. Flora, 185, 107-125.
[14]	Lasa B, Frechilla S, Aparicio-Tejo PM, Lamsfus C (2002). Role of glutamate dehydrogenase and phosphoenol- pyruvate carboxylase activity in ammonium nutrition tolerance in roots. Plant Physiology and Biochemistry, 40, 969-976.
[15]	Levizou E, Petropoulou Y, Manetas Y (2004). Carotenoid composition of peridermal twigs does not fully conform to a shade acclimation hypothesis. Photosynthetica, 42, 591-596.
[16]	Lichtenthaler HK (1987). Chorophylls and carotenoids: Pigments of photosynthetic biomembranes. Methods in Enzymology, 148, 350-382.
[17]	Nilsen ET (1995). Stem photosynthesis: extent, patterns and role in plant carbon economy. In: Garter B ed. Plant Stems: Physiology and Functional Morphology. Academic Press, San Diego. 223-240.
[18]	Pfanz H (2008). Bark photosynthesis. Trees, 22, 137-138.
[19]	Pfanz H, Aschan G (2001). The existence of bark and stem photosynthesis in woody plants and its significance for the overall carbon gain. An eco-physiological and ecological approach. Botany, 62, 478-510.
[20]	Rosendahl L, Vance CP, Pedersen WB (1990). Products of dark CO₂ fixation in pea root nodules support bacteroid metabolism. Plant Physiology, 93, 12-19. URL PMID
[21]	Shi JM (施建敏), Guo QR (郭起荣), Yang GY (杨光耀) (2005). Study on the photosynthetic dynamic variation of Phyllostachys edulis. Forest Research (林业科学研究), 18, 551-555. (in Chinese with English abstract)
[22]	Tokarz K, Pilarski J (2005). Optical properties and the content of photosynthetic pigments in the stems and leaves of the apple-tree. Acta Physiologiae Plantarum, 27, 183-191.
[23]	Valenini R, Matteucci G, Dolman AJ, Schulze ED, Rebmann C, Moors EJ, Granier A, Gross P, Jensen NO, Pilegaard K, Lindroth A, Grelle A, Bernhofer C, Grllnwald T, Aubinet M, Ceulemans R, Kowalski AS, Vesala1 T, Rannik1 U, Berbigier P, Loustau D, Gumundsson J, Thorgeirsson H, Ibrom A, Morgenstern K, Clement R, Moncrieff J, Montagnani L, Minerbi S, Jarvis PG (2000). Respiration as the main determinant of carbon balance in European forests. Nature, 404, 861-865. DOI URL PMID
[24]	Wang WJ (王文杰), Zhang ZH (张衷华), Zu YG (祖元刚), He HS (贺海升), Guan Y (关宇), Li WX (李文馨) (2009). Photosynthetic characteristics of the non-photosynthetic organs of Mikania micrantha and its ecological significance. Acta Ecologica Sinica (生态学报), 29, 28-36. (in Chinese with English abstract)
[25]	Wang WJ (王文杰), Zu YG (祖元刚), Wang HM (王慧梅) (2007). Review on the photosynthetic function of non- photosynthetic woody organs of stem and branches. Acta Ecologica Sinica (生态学报), 27, 1583-1595. (in Chinese with English abstract)
[26]	Wang WJ, Watanabe Y, Endo I, Kitaoka S, Koike T (2006a). Seasonal changes in the photosynthetic capacity of cones on a larch (Larix kaempferi) canopy. Photosynthetica, 44, 345-348.
[27]	Wang WJ, Zu YG, Cui S, Hirano T, Watanabe Y, Koike T (2006 b). Carbon dioxide exchange of larch (Larix gmelinii) cones during development. Tree Physiology, 26, 1363-1368. DOI URL PMID
[28]	Wilson DR, Martinez TR (1997). Improved heterogeneous distance functions. Journal of Artificial Intelligence Research, 6, 1-34.
[29]	Wittmann C, Aschan G, Pfanz H (2001). Leaf and twig photosynthesis of young beech (Fagus sylvatica) and aspen(Populus tremula) trees grown under different light regime. Basic and Applied Ecology, 2, 145-154.
[30]	Wittmann C, Pfanz H, Loreto F, Centritto M, Pietrini F, Alessio G (2006). Stem CO₂ release under illumination: corticular photosynthesis, photorespiration or inhibition of mitochondrial respiration? Plant, Cell & Environment, 29, 1149-1158. URL PMID
[31]	Zu YG (祖元刚), Zhang ZH (张衷华), Wang WJ (王文杰), Yang FJ (杨逢建), He HS (贺海升) (2006). Different characteristics of photosynthesis in stems and leaves of Mikania micranth. Journal of Plant Ecology (Chinese Version) (植物生态学报), 30, 998-1004. (in Chinese with English abstract)