Effects of Shading on Photosynthesis and Aromatics of Pandan (Pandanus amaryllifolius) Plants
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摘要:
目的 研究不同荫蔽度对香露兜光合特性、生长情况及香气成分的影响,为生产上林下复合种植香露兜提供理论依据。 方法 采用人工荫蔽盆栽试验的方法,设置全光照、30%荫蔽度、60%荫蔽度和90%荫蔽度等4种不同荫蔽处理,探究不同荫蔽度处理下香露兜光合参数、生长指标和香气成分对光环境的响应。 结果 荫蔽度为30%和60%时,香露兜净光合速率和气孔导度均显著高于全光照和90%遮荫处理,有利于提高香露兜的光合作用,植株叶片较多。随着荫蔽度的增加,分蘖数显著减少。4种遮荫处理下香露兜共鉴定出27种挥发性物质,2-乙酰-1-吡咯啉、叶绿醇、角鲨烯、丙醇、丙酮醇、3-甲基-2-(5H)-呋喃酮、新植二烯、棕榈酸乙酯、2,3-二氢苯并呋喃和亚油酸乙酯等10种共有香气成分含量差异显著。30%和60%荫蔽处理下香露兜叶片关键特征香气物质2-乙酰-1-吡咯啉和3-甲基-2-(5H)-呋喃酮、2,3-二氢苯并呋喃、亚油酸乙酯和叶绿醇显著高于其他处理,30%荫蔽处理角鲨烯和叶绿醇含量显著高于其他处理,60%荫蔽处理丙酮醇、2-乙酰-1-吡咯啉、3-甲基-2-(5H)-呋喃酮、新植二烯、棕榈酸乙酯和亚油酸乙酯显著高于其他处理。 结论 30%~60%荫蔽度可促进香露兜生长,提高主要香气成分含量,风味品质佳。 Abstract:Objective Effects of shading on photosynthesis, growth, and aromatic composition of Pandan (Pandanus amaryllifolius) plants were studied for cultivation improvement. Method In a pot experimentation, pandan plants were grown under either full sun exposure or 30%, 60% or 90% shading. The photosynthesis, growth, and aromatics of the plants were monitored. Result The net photosynthetic rate, stomatal conductance, and count of the leaves on plants grown under 30% and 60% shading were significantly higher than those under either full sun exposure or 90% shading. On the other hand, the tiller number decreased significantly with increasing shading. Twenty-seven volatile aromatic compounds were identified regardless the difference in treatment. There were 10 aromatics, including 2-acetyl-1-pyrrolidine phytol, squalene, 1-propanol, acetol, 3-methyl-2-(5H)-furanone, neophytadiene, ethyl palmitate, 2,3-dihydrobenzofuran, and ethyl linoleate, commonly present, but differed significantly in relative content when the plants were under the varied shading. For instance, the contents of 2-acetyl-1-pyrroline, 3-methyl-2-(5H)-furanone, 2,3-dihydrobenzofuran, ethyl linoleate, and phytol were significantly higher under 30% and 60% shading, those of squalene and phytol significantly higher under 30% shading, and those of acetol, 2-acetyl-1-pyrroline, 3-methyl-2-(5H)-furanone, neophytadiene, ethyl palmitate, and ethyl linoleate significantly higher under 60% shading than the other treatments. Conclusion By artificially imposing 30% or 60% shading, pandan plants grew more vigorously generating more fragrant substances in the leaves than exposing naturally to the sun. -
Key words:
- Pandanus amaryllifolius /
- shading /
- photosynthetic characteristics /
- aroma components
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图 3 遮荫处理下香露兜叶片香气成分种类数量(A)和含量(B)
注:图中不同小写字母表示4种遮荫处理间同种香气成分化合物种类的差异显著性水平(P<0.05);Ⅰ-吡咯类,Ⅱ-醇类,Ⅲ-呋喃类,Ⅳ-酚类,Ⅴ-呋喃酮类,Ⅵ-烃类,Ⅶ-酮类,Ⅷ-酯类.
Figure 3. Categories (A) and contents (B) of volatile compounds in leaves of pandan plant grown under varied shading treatments
Note: Data with different lowercase letters indicate significant difference on same aromatic compound among treatments at 0.05 level; Ⅰ-Pyrroles,Ⅱ-Alcohols,Ⅲ-Furans,Ⅳ-Phenols,Ⅴ-Furanones,Ⅵ-Hydrocarbons,Ⅶ-Ketones,Ⅷ-Esters.
图 4 不同荫蔽度香露兜叶片香气成分PCA分析
注:A.香气成分组成主成分分析;B.香气成分种类主成分分析;代码为香气成分(表1);黑色圆点表示香露兜27种香气成分,黑色三角表示香露兜8种香气成分种类。
Figure 4. Biplot of principal components analysis showing aromatics in leaves of pandan plants grown under varied shading treatments
Note: A: on aromatic compounds; B: on categories of aromatics; sample codes: for aromatic components (Table 1); black dots: 27 aromatic compounds; black triangles: 8 classes of aromatics in pandan plants.
表 1 遮荫处理下香露兜叶片挥发性香气成分及含量
Table 1. Volatiles in leaves of pandan plant grown under varied shading treatments
化合物种类
classes保留时间
RT保留指数
LRI化合物名称
compounds代码
code遮荫度 The shade degree/% 0 30 60 90 脂类 Esters 9.67 1217 丙酮酸甲酯 methyl pyruvate H1 12.58±0.37 8.23±0.45 ― ― 12.07 1349 L-乳酸乙酯(−)-Ethyl L-lactate H2 31.07±1.04 11.47±0.33 ― 4.88±0.12 14.24 1436 乙醇酸乙酯 Ethyl 2-hydroxyacetate H3 8.36±0.22 2.30±0.09 2.92±0.05 ― 15.21 1474 (S)-缩水甘油乙酸酯
oxiran-2-ylmethyl acetateH4 ― 1.33±0.02 ― ― 29.93 2142 (±)-α-羟基-γ-丁内酯 α-butyrolactone H5 24.76±0.11 19.82±0.25 33.56±0.15 ― 31.65 2251 棕榈酸乙酯 ethyl palmitate H6 21.73±0.19 1.55±0.05 34.44±0.70 14.00±0.27 34.86 2471 油酸乙酯 ethyl oleate H7 ― 5.30±0.06 7.24±0.29 3.32±0.19 35.49 2521 亚油酸乙酯 Ethyl linoleate H8 12.73±0.78 25.71±0.51 37.54±1.62 19.78±0.48 36.59 2591 亚麻酸乙酯 ethyl linolenate H9 22.91±0.96 ― ― ― 醇类 Alcohols 7.62 1036 丙醇 1-propanol B1 4.02±0.04 3.78±0.05 3.77±0.16 4.29±0.15 9.44 1209 异戊醇 isoamyl alcohol B2 2.27±0.08 1.16±0.10 ― ― 20.73 1658 1,2-乙二醇 1,2-ethanediol B3 13.95±0.68 ― ― ― 36.82 2622 叶绿醇 Phytol B4 62.40±1.28 149.23±0.64 124.37±0.59 70.63±0.96 烃类 Hydrocarbons 13.14 1400 正十四烷 tetradecane F1 3.12±0.01 ― ― ― 19.08 1600 正十六烷 hexadecane F2 6.70±1.15 3.13±0.21 ― ― 23.66 1800 十八烷 octadecane F3 ― 1.33±0.01 ― ― 25.99 1922 新植二烯 neophytadiene F4 16.05±0.21 26.36±0.26 54.67±0.87 36.66±0.18 39.85 2865 角鲨烯 Squalene F5 609.38±4.09 784.72±6.87 413.47±17.74 41.64±1.5 酮类 Ketones 10.83 1284 3-羟基-2-丁酮 acetoin G1 9.38±0.51 5.11±0.03 9.02±0.18 ― 11.23 1303 丙酮醇 acetol G2 95.15±5.99 87.36±0.51 155.76±0.9 20.01±0.19 20.80 1668 1,2-环己二酮 Cyclohexan-1,2-dion G3 ― ― 4.37±0.09 ― 24.25 1830 甲基环戊烯醇酮 cyclotene G4 ― ― 3.21±0.64 ― 25.49 1894 3-甲基环戊烷-1,2-二酮
3-ethyl-1,2-cyclopentanedioneG5 ― ― 3.16±0.06 ― 吡咯类 Pyrroles 11.79 1330 2-乙酰基-1-吡咯啉 2-acetyl-1-pyrrolidine A1 5.41±0.08 11.76±0.21 13.88±1.64 4.48±0.14 酚类 Phenols 32.62 2318 2,4-二叔丁基苯酚 2,4-di-tert-butylphenol C1 12.97±0.12 15.16±0.43 ― ― 呋喃 Furans 33.66 2389 2,3-二氢苯并呋喃 coumaran D1 61.94±5.80 86.00±7.22 94.92±14.73 19.82±1.13 呋喃酮 Furanone 21.64 1713 3-甲基-2-(5H)-呋喃酮
3-methyl-2(5H)-furanoneE1 32.04±0.44 43.92±1.03 55.46±0.17 16.27±0.11 注:“―”表示未检测到该物质;数据后不同小写字母表示4种遮荫处理间香气成分含量的差异显著性水平(P<0.05)。
Note: “―” indicates substance not detected; data with different lowercase letters indicate significant difference on aromatic contents among treatments at 0.05 level. -
[1] 孙祥钟. 中国植物志: 第八卷[M]. 北京: 科学出版社, 1992: 12-23. [2] ROUTRAY W, RAYAGURU K. Chemical constituents and post-harvest prospects of Pandanus amaryllifolius leaves: a review [J]. Food Reviews International, 2010, 26(3): 230−245. doi: 10.1080/87559129.2010.484114 [3] PETER K V. Handbook of Herbs and Spices[M], Elsevier, 2012. [4] 尹桂豪, 王明月, 曾会才. 香露兜叶挥发油的超临界萃取及气相色谱-质谱联用分析 [J]. 时珍国医国药, 2010, 21(1):159−160. doi: 10.3969/j.issn.1008-0805.2010.01.078YIN G H, WANG M Y, ZENG H C. Supercritical extraction and GC-MS analysis of volatile oil from Pandan leaves [J]. Lishizhen Medicine and Materia Medica Research, 2010, 21(1): 159−160.(in Chinese) doi: 10.3969/j.issn.1008-0805.2010.01.078 [5] CHONG H Z, ASMAH R, MD A A, et al. Chemical analysis of pandan leaves (Pandanus amaryllifolius) [J]. International Journal of Natural Product and Pharmaceutical Sciences, 2010, 1(1): 7−10. [6] JIANG J. Volatile composition of pandan leaves (Pandanus amaryllifolius) [J]. Flavor Chemistry of Ethnic Foods, 1999: 105−109. [7] MAR A, MAR A A, THIN P P, et al. Study on the phytochemical constituents in essential oil of Pandanus amaryllifolius Roxb. leaves and their anti-bacterial efficacy [J]. Yadanabon University Research Journal, 2019, 101(1): 1−3. [8] WAKTE K V, THENGANE R J, JAWALI N, et al. Optimization of HS-SPME conditions for quantification of 2-acetyl-1-pyrroline and study of other volatiles in Pandanus amaryllifolius Roxb [J]. Food Chemistry, 2010, 121(2): 595−600. doi: 10.1016/j.foodchem.2009.12.056 [9] SUROJANAMETAKUL V, BOONBUMRUNG S, TUNGTRAKUL P, et al. Encapsulation of natural flavor from Pandanus amaryllifolius roxb. in rice starch aggregates [J]. Food Science and Technology Research, 2019, 25(4): 577−585. doi: 10.3136/fstr.25.577 [10] LAKSANALAMAI V, ILANGANTILEKE S. Comparison of aroma compound (2-acetyl-1-pyrroline) in leaves from pandan (Pandanus amaryllifolius) and Thai fragrant rice (Kaho Dwak Mali-105) [J]. Cereal Chemistry, 1993, 70(4): 381−384. [11] 程亚娇, 范元芳, 谌俊旭, 等. 光照强度对大豆叶片光合特性及同化物的影响 [J]. 作物学报, 2018, 44(12):1867−1874. doi: 10.3724/SP.J.1006.2018.01867CHENG Y J, FAN Y F, SHEN J X, et al. Effects of light intensity on photosynthetic characteristics and assimilates of soybean leaf [J]. Acta Agronomica Sinica, 2018, 44(12): 1867−1874.(in Chinese) doi: 10.3724/SP.J.1006.2018.01867 [12] 庄辉发, 王辉, 王华, 等. 不同荫蔽度对香草兰光合特性的影响 [J]. 热带农业科学, 2013, 33(10):16−18. doi: 10.3969/j.issn.1009-2196.2013.10.004ZHUANG H F, WANG H, et al. Effect of different shade degrees on Vanilla photosynthetic characteristics [J]. Chinese Journal of Tropical Agriculture, 2013, 33(10): 16−18.(in Chinese) doi: 10.3969/j.issn.1009-2196.2013.10.004 [13] 庄辉发, 王辉, 王华, 等. 不同荫蔽度对香草兰光合作用与产量的影响 [J]. 江苏农业科学, 2012, 40(8):239−240. doi: 10.3969/j.issn.1002-1302.2012.08.095ZHUANG H F, WANG H, Wang H, et al. Effect of different shade degrees on Vanilla photosynthetic characteristics and yield [J]. Jiangsu Agricultural Sciences, 2012, 40(8): 239−240.(in Chinese) doi: 10.3969/j.issn.1002-1302.2012.08.095 [14] 赵思毅, 黄承建, 肖万林, 等. 不同程度荫蔽胁迫对苎麻光合特性的影响 [J]. 中国麻业科学, 2017, 39(1):19−24. doi: 10.3969/j.issn.1671-3532.2017.01.004ZHAO S Y, HUANG C J, XIAO W L, et al. Effects of varying shade levels on photosynthetic characteristics of ramie [J]. Plant Fibers and Products, 2017, 39(1): 19−24.(in Chinese) doi: 10.3969/j.issn.1671-3532.2017.01.004 [15] 汤永坚, 汪益磊, 肖枫, 等. 收获前15d持续遮光对香稻产量、品质和香气的影响 [J]. 西南农业学报, 2015, 28(3):939−943.TANG Y J, WANG Y L, XIAO F, et al. Effect of continuous shading treatment 15 days before harvest on grain yield, quality and aroma of aromatic rice [J]. Southwest China Journal of Agricultural Sciences, 2015, 28(3): 939−943.(in Chinese) [16] 沈生荣, 杨贤强, 陈席卿. 遮荫对蒸青绿茶香气成分的影响 [J]. 浙江农业大学学报, 1990(1):98−101.SHEN S R, YANG X Q, CHEN X Q. Effect of shading on the flavor characters of steam-green-tea [J]. Journal of Zhejiang University, 1990(1): 98−101.(in Chinese) [17] VOLPICELLI G, BOERO E, SVERZELLATI N, et al. Semi-quantification of pneumothorax volume by lung ultrasound [J]. Intensive Care Medicine, 2014, 40(10): 1460−1467. doi: 10.1007/s00134-014-3402-9 [18] LOH S K, CHE MAN Y B, TAN C P, et al. Process optimisation of encapsulated pandan (Pandanus amaryllifolius) powder using spray-drying method [J]. Journal of the Science of Food and Agriculture, 2005, 85(12): 1999−2004. doi: 10.1002/jsfa.2169 [19] 李冬林, 金雅琴, 崔梦凡, 等. 夏季遮光对连香树幼苗形态、光合作用及叶肉细胞超微结构的影响 [J]. 浙江农林大学学报, 2020, 37(3):1−10. doi: 10.11833/j.issn.2095-0756.20190396LI D L, JIN Y Q, CUI M F, et al. Growth, photosynthesis and ultrastructure of mesophyll cells for Cercidiphyllum japonicum seedlings with shading in summer [J]. Journal of Zhejiang A&F University, 2020, 37(3): 1−10.(in Chinese) doi: 10.11833/j.issn.2095-0756.20190396 [20] 薛黎, 李志辉, 童方平, 等. 遮荫对闽楠幼苗光合及其叶片解剖特性的影响 [J]. 西北植物学报, 2019, 39(7):1221−1229.XUE L, LI Z H, TONG F P, et al. Effect of shading on photosynthetic characteristics and anatomical structure in leaves of Phoebe bournei seedlings [J]. Acta Botanica Boreali-Occidentalia Sinica, 2019, 39(7): 1221−1229.(in Chinese) [21] 郑坚, 吴朝辉, 陈秋夏, 等. 遮荫对降香黄檀幼苗生长和生理的影响 [J]. 林业科学, 2016, 52(12):50−57. doi: 10.11707/j.1001-7488.20161206ZHENG J, WU Z H, CHEN Q X, et al. Influence of shading on growth and physiology of Dalbergia odorifera seedlings [J]. Scientia Silvae Sinicae, 2016, 52(12): 50−57.(in Chinese) doi: 10.11707/j.1001-7488.20161206 [22] 刘贤赵, 唐绍忠. 番茄不同生育阶段遮荫对光合作用与产量的影响 [J]. 园艺学报, 2002, 29(5):427−432. doi: 10.3321/j.issn:0513-353X.2002.05.006LIU X Z, AND K. Effects of shading on photosynthesis and yield of tomato plants at different growth stages [J]. Acta Horticulturae Sinica, 2002, 29(5): 427−432.(in Chinese) doi: 10.3321/j.issn:0513-353X.2002.05.006 [23] WANG M, SHI S, LIN F, et al. Effects of soil water and nitrogen on growth and photosynthetic response of Manchurian ash (Fraxinus mandshurica) seedlings in northeastern China [J]. PLoS One, 2012, 7(2): e30754. doi: 10.1371/journal.pone.0030754 [24] MONNEVEUX P, PASTENES C, REYNOLDS M P. Limitations to photosynthesis under light and heat stress in three high yielding wheat genotypes [J]. Journal of Plant Physiology, 2003, 160(6): 657−666. doi: 10.1078/0176-1617-00772 [25] Zhu J J, XU M. Impact of the Shading Intensity on the Cultivation Characteristics of Phlox subulata [J]. Botanical Research, 2017, 6(5): 298−303. doi: 10.12677/BR.2017.65039 [26] 胡卫丽, 朱旭, 杨厚勇, 等. 荫蔽胁迫对不同绿豆品种生物学性状及产量的影响 [J]. 安徽农业科学, 2017, 45(1):60−63, 75. doi: 10.3969/j.issn.0517-6611.2017.01.019HU W L, ZHU X, YANG H Y, et al. Effects of shading stress on the biological traits and yield of different Mung bean varieties [J]. Journal of Anhui Agricultural Sinica, 2017, 45(1): 60−63, 75.(in Chinese) doi: 10.3969/j.issn.0517-6611.2017.01.019 [27] 于晓波, 梁建秋, 何泽民, 等. 玉米-大豆带状套作对大豆叶片形态及光合特性的影响 [J]. 中国油料作物学报, 2016, 38(4):452−459. doi: 10.7505/j.issn.1007-9084.2016.04.007YU X B, LIANG J Q, HE Z M, et al. Response of leaf morphology and photosynthetic characteristics of soybean in maize-soybean relay strip intercropping system [J]. Chinese Journal of Oil Crop Sciences, 2016, 38(4): 452−459.(in Chinese) doi: 10.7505/j.issn.1007-9084.2016.04.007 [28] 张惠君, 林海波, 马野夫, 等. 耐密植大豆品种沈农12叶片性状研究 [J]. 大豆科学, 2010, 29(2):233−237.ZHANG H J, LIN H B, MA Y F, et al. Leaf traits of soybean cultivar Shennong 12 with tolerance to high planting density [J]. Soybean Science, 2010, 29(2): 233−237.(in Chinese) [29] LAOHAKUNJIT N, NOOMHORM A. Supercritical carbon dioxide extraction of 2-acetyl-1-pyrroline and volatile components from pandan leaves [J]. Flavour and Fragrance Journal, 2004, 19(3): 251−259. doi: 10.1002/ffj.1297 [30] WAKTE K V, ZANAN R L, THENGANE R J, et al. Identification of elite population of Pandanus amaryllifolius roxb. for higher 2-acetyl-1-pyrroline and other volatile contents by HS-SPME/GC-FID from peninsular India [J]. Food Analytical Methods, 2012, 5(6): 1276−1288. doi: 10.1007/s12161-012-9373-y [31] DAS B, SENGUPTA S, KOLE B, et al. Elucidation of aroma levels within a set of rice landraces by means of aroma linked markers [J]. Journal of Pharmacognosy and Phytochemistry, 2018, 7(4): 1756− 1760. [32] SILALAHI M, SI M. Pandanus amaryllifolius Roxb(Pemanfaatan dan potensinya sebagai pengawet makanan) [J]. Pro-Life, 2018, 5(3): 626−636. doi: 10.33541/pro-life.v5i3.842 [33] SCHILLING M W, PHAM-MONDALA A J, DHOWLAGHAR N, et al. Changes in the volatile composition of fresh pork sausage with natural antioxidants during long-term frozen storage [J]. Meat and Muscle Biology, 2019, 3(1): 194. doi: 10.22175/mmb2019.03.0007 [34] SOMTA P, KUSWANTO K, SRINIVES P. The genetics of pandan-like fragrance, 2-acetyl-1-pyrroline, in crops [J]. Agrivita : Journal of Agricultural Science, 2019, 41(1): 10−22. [35] SCHIEBERLE P. The role of free amino acids present in yeast as precursors of the odorants 2-acetyl-1-pyrroline and 2-acetyltetrahydropyridine in wheat bread crust [J]. Zeitschrift Für Lebensmittel-Untersuchung Und -Forschung, 1990, 191(3): 206−209. doi: 10.1007/BF01197621 [36] YOSHIHASHI T, HUONG N T T, INATOMI H. Precursors of 2-acetyl-1-pyrroline, a potent flavor compound of an aromatic rice variety [J]. Journal of Agricultural and Food Chemistry, 2002, 50(7): 2001−2004. doi: 10.1021/jf011268s [37] 莫钊文, 范平珊, 潘圣刚, 等. 肥料类型及施用方式对香稻香气2-乙酰-1-吡咯啉含量的影响 [J]. 华北农学报, 2016, 31(5):152−158. doi: 10.7668/hbnxb.2016.05.023MO Z W, FAN P S, PAN S G, et al. Effect of fertilizer types and fertilization methods on 2-acetyl-1-pyrroline content in aromatic rice [J]. Acta Agriculturae Boreali-Sinica, 2016, 31(5): 152−158.(in Chinese) doi: 10.7668/hbnxb.2016.05.023