南京林业大学学报(自然科学版) ›› 2023, Vol. 47 ›› Issue (1): 109-120.doi: 10.12302/j.issn.1000-2006.202109027
收稿日期:
2021-09-14
接受日期:
2021-11-27
出版日期:
2023-01-30
发布日期:
2023-02-01
通讯作者:
王曙光
基金资助:
JIN Diankun(), LYU Zhuo, WANG Shuguang(), LONG Hao, ZHANG Chong, WANG Shenghan
Received:
2021-09-14
Accepted:
2021-11-27
Online:
2023-01-30
Published:
2023-02-01
Contact:
WANG Shuguang
摘要: 【目的】 探究竹子营养叶与秆箨之间的关系,揭示竹子不同功能叶器官的结构差异,为竹类植物的基础生物学研究提供新的理论信息。【方法】 以6种竹子即勃氏甜龙竹(Dendrocalamus brandisii)、慈竹(Bambusa emeiensis )、绵竹(Lingnania intermedia)、香竹(Chimonocalamus delicatus)、云南箭竹(Fargesia yunnanensis)和美竹(Phyllostachys mannii)的营养叶、秆箨和叶枝为研究对象,观察它们的显微结构,并对其各项指标进行测量与比较。【结果】 竹子营养叶解剖结构具有指状臂细胞或梅花状细胞的分化,并且具有薄壁维管鞘细胞;秆上部的箨片形态更接近于叶片,而下部的箨片具有较厚的角质层,无叶肉细胞的分化,与叶片形态差异明显。竹子营养叶的叶片、叶柄、叶鞘和叶枝维管束韧皮部的面积占比均高于木质部,与秆箨(笋箨)有显著差异。秆箨(笋箨)的箨鞘与箨片维管束木质部面积占比更高,虽然秆上部箨片形状和大小都与叶片接近,但与叶片相比,其维管束木质部面积占比更高。叶鞘和箨鞘的解剖结构与叶片和箨片的有区别,与叶柄的结构基本一致,且基部包裹着枝或腋芽。【结论】 竹子营养叶器官韧皮部更发达,秆箨木质部更发达,箨鞘应属变态叶柄。竹类植物所有器官或组织的韧皮部均分布在维管束的远轴面,木质部则分布在维管束的近轴面。在竹子的叶器官中,营养叶以及叶枝主要以光合同化物运输为主,而秆箨在生理功能上除了机械支持作用,更多的是完成水分运输。
中图分类号:
金点坤,吕卓,王曙光,等. 6种竹子叶器官的解剖结构比较[J]. 南京林业大学学报(自然科学版), 2023, 47(1): 109-120.
JIN Diankun, LYU Zhuo, WANG Shuguang, LONG Hao, ZHANG Chong, WANG Shenghan. Comparison of anatomical structure of six bamboo species cotyledon organs[J].Journal of Nanjing Forestry University (Natural Science Edition), 2023, 47(1): 109-120.DOI: 10.12302/j.issn.1000-2006.202109027.
表1
6种竹子营养叶、叶鞘及枝条解剖特征"
指标 index | 勃氏甜龙竹 D. brandisii | 慈竹 B. emeiensis | 绵竹 L. intermedia | 香竹 C. delicatus | 云南箭竹 F. yunnanensis | 美竹 P. mannii | ||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
叶长/cm leaf length | 28.08±3.76 a | 20.90±2.88 b | 9.87±0.88 e | 8.76±1.57 e | 17.52±2.44 c | 15.29±2.11 d | ||||||||
叶宽/cm leaf width | 4.67±1.24 a | 2.30±0.41 b | 1.23±0.34 d | 1.23±0.16 d | 1.81±0.34 c | 1.93±0.32 c | ||||||||
长宽比aspect ratio | 6.01 e | 9.08 b | 8.02 c | 7.12 d | 9.67 a | 7.92 c | ||||||||
中部厚度/μm centre thickness | 叶片leaf blade | 104.41±7.99 Gb | 90.84±11.78 Fd | 97.35±4.92 Fc | 107.04±10.25 Eb | 81.19±8.64 Ge | 124.56±12.34 Ga | |||||||
叶柄petiole | 599.93±28.61 Aa | 360.32±3.37 Ad | 325.97±13.65 Ce | 463.46±9.56 Ab | 443.55±13.42 Bc | 357.63±3.78 Bd | ||||||||
叶鞘leaf sheath | 264.39±15.68 Ec | 150.45±6.92 Ed | 340.12±20.74 BCb | 105.42±16.83 Ef | 141.94±6.68 Fe | 364.97±20.02 Ba | ||||||||
叶枝twig | 447.81±63.96 Ba | 272.17±4.40 Cbc | 290.77±61.80 Db | 257.56±18.26 Bcd | 239.89±15.01 Ede | 225.73±2.08 De | ||||||||
秆上部箨片upper culm blade | 158.84±19.11 Fc | 148.85±23.01 Ed | 262.15±14.90 Ea | 190.04±14.04 CDb | 129.32±20.25 Fe | 144.16±6.57 Fd | ||||||||
秆下部箨片lower culm blade | 344.40±36.11 Db | 183.33±23.63 De | 329.60±4.59 Cc | 171.65±9.44 Df | 775.14±4.77 Aa | 267.76±3.16 Cd | ||||||||
秆上部箨鞘upper culm sheath | 459.59±8.14 Ba | 189.48±14.66 Dd | 360.05±12.70 Bb | 181.98±31.32 Dde | 317.64±6.26 Dc | 178.67±15.43 Ee | ||||||||
秆下部箨鞘lower culm sheath | 374.81±33.40 Cb | 327.02±8.09 Bc | 488.29±17.77 Aa | 206.28±3.36 Cd | 379.01±5.48 Cb | 381.10±2.39 Ab | ||||||||
近轴面表皮 厚度/μm adaxial epidermis thickness | 叶片leaf blade | 12.60±2.58 Bb | 10.32±2.28 Cc | 13.84±2.64 Ba | 9.26±1.69 Cd | 10.99±1.62 Bc | 8.01±0.98 De | |||||||
叶柄petiole | 16.46±1.98 Aa | 13.17±2.09 Bb | 11.28±1.67 CDc | 11.15±1.98 Bc | 10.72±2.07 Bc | 8.99±1.08 Cd | ||||||||
叶鞘leaf sheath | 7.36±1.32 Db | 5.75±1.46 Fc | 8.14±1.82 Ea | 5.08±1.00 Ec | 5.74±1.50 DEc | 5.57±1.67 Fc | ||||||||
秆上部箨片upper culm blade | 12.15±2.62 Bc | 14.26±3.39 Ab | 12.62±6.60 Cbc | 16.33±3.62 Aa | 11.37±1.99 Bcd | 9.86±2.22 Bd | ||||||||
秆下部箨片lower culm blade | 12.09±2.85 Bc | 9.45±2.08 Ce | 15.28±3.69 Aa | 11.25±3.25 Bdc | 13.59±2.75 Ab | 10.42±3.55 Ace | ||||||||
秆上部箨鞘upper culm sheath | 4.93±1.12 Ed | 7.87±1.12 Ea | 6.59±1.60 Fc | 5.34±1.02 Ed | 6.27±1.53 Dc | 7.02±1.30 Eb | ||||||||
秆下部箨鞘lower culm sheath | 9.91±1.95 Ca | 9.96±1.61 Ca | 10.24±2.31 Da | 6.04±1.97 Dc | 6.86±1.93 Dbc | 7.32±1.50 Eb | ||||||||
远轴面表皮 厚度/μm abaxial epidermis thickness | 叶片leaf blade | 10.14±1.48 Ca | 7.61±1.43 Ed | 9.33±1.57 Cb | 8.55±1.67 Cc | 8.04±0.68 Ecd | 7.74±0.92 Dd | |||||||
叶柄petiole | 8.68±1.26 Db | 9.23±1.60 Cab | 9.54±2.28 Ca | 8.46±2.40 Cb | 7.06±1.19 Fc | 7.49±0.97 Dc | ||||||||
叶鞘leaf sheath | 9.15±0.99 Dc | 9.32±1.32 Cbc | 10.28±1.90 Bab | 8.39±1.99 Cc | 10.54±3.00 Da | 9.36±2.43 Cbc | ||||||||
叶枝twig | 7.47±1.04 Eb | 5.81±1.53 Fc | 6.04±1.14 Dc | 6.19±1.40 Ec | 12.06±1.94 BCa | 6.26±0.93 Ec | ||||||||
秆上部箨片upper culm blade | 4.83±1.50 Fe | 7.73±2.07 Ec | 9.90±1.85 BCa | 8.60±1.73 Cb | 8.52±1.45 Ebc | 5.99±1.20 Ed | ||||||||
秆下部箨片lower culm blade | 7.12±1.72 Ed | 8.40±1.67 Dc | 9.87±2.26 BCb | 7.41±1.51 Dd | 11.78±3.41 Ca | 9.86±1.66 BCb | ||||||||
秆上部箨鞘upper culm sheath | 13.12±2.09 Bd | 12.43±1.91 Ad | 9.84±1.76 BCe | 25.51±4.17 Aa | 17.74±2.49 Ab | 15.21±3.39 Ac | ||||||||
秆下部箨鞘lower culm sheath | 16.13±2.64 Aa | 11.52±1.50 Bc | 13.15±2.11 Ab | 16.23±2.39 Ba | 12.94±3.69 Bb | 10.02±1.69 Bd | ||||||||
维管束 面积/μm2 vascular bundle area | 叶片leaf blade | 2 530.76±320.82 Gb | 2 364.15±196.90 Fc | 2 122.83±136.13 Fd | 2 846.46±491.35 Ga | 2 502.30±341.81 Hbc | 2 507.99±162.96 Gbc | |||||||
叶柄petiole | 2 781.78±441.93 Gb | 1 670.84±197.54 Gd | 2 268.06±232.61 Fc | 3 111.31±445.91 Ga | 2 834.18±205.98 Gb | 2 245.66±226.00 Gc | ||||||||
叶鞘leaf sheath | 7 514.30±974.40 Fa | 3 870.86±477.88 Ed | 3 262.26±469.15 Ee | 4 119.75±574.76 Ed | 5 689.84±401.51 Ec | 6 146.79±542.80 Eb | ||||||||
叶枝twig | 12 970.38±2 044.21 Da | 4 466.00±393.08 Db | 4 986.64±907.23 Db | 3 811.19±465.81 Fb | 4 067.46±392.10 Fb | 4 654.33±415.77 Fb | ||||||||
秆上部箨片upper culm blade | 15 273.25±1 223.59 Cc | 17 675.32±1 884.23 Cb | 22 345.13±408.04 Ba | 7 240.21±861.27 De | 10 813.09±597.18 Dd | 7 747.71±497.77 De | ||||||||
秆下部箨片lower culm blade | 42 228.37±6 069.38 Aa | 19 906.20±2 018.07 Ac | 22 978.22±3 910.28 Bb | 15 105.35±1 636.78 Ad | 41 401.81±3 922.77 Aa | 8 175.83±886.04 Ce | ||||||||
秆上部箨鞘upper culm sheath | 11 619.41±1 393.04 Ed | 19 031.83±1 387.74 Bb | 26 370.91±1 587.41Aa | 8 227.78±1 308.61 Cf | 12 499.79±996.84 Cc | 10 661.93±1 108.18 Ae | ||||||||
秆下部箨鞘lower culm sheath | 19 663.36±3 347.66 Bb | 20 010.24±2 702.30 Aab | 20 649.12±2 840.97 Ca | 11 661.58±1 216.91 Bd | 16 854.24±2 690.98 Bc | 8 967.64±1 075.82 Be | ||||||||
木质部 面积/μm2 xylem area | 叶片leaf blade | 382.94±42.06 Fa | 263.87±29.43 Gc | 205.06±14.33 Fd | 323.98±123.63 Gb | 389.09±111.24 Fa | 386.75±49.13 Fa | |||||||
叶柄petiole | 242.93±33.36 Gd | 147.91±22.21 He | 238.23±31.22 Fd | 409.00±49.37 Fa | 341.00±46.57 Fb | 317.82±34.56 Gc | ||||||||
叶鞘leaf sheath | 1 617.51±142.80 Ea | 572.41±77.64 Fd | 503.02±71.43 Ee | 412.50±80.28 Ff | 742.86±58.82 Ec | 792.44±62.77 Db | ||||||||
叶枝twig | 1 600.11±347.49 Ea | 690.00±92.99 Ec | 594.61±110.00 Dd | 809.22±67.44 Eb | 705.95±63.99 Ec | 523.95±89.52 Ed | ||||||||
秆上部箨片upper culm blade | 15 273.25±1 223.59 Ac | 17 675.32±1 884.23 Ab | 22 345.13±408.04 Aa | 7 240.21±861.27 Ae | 10 813.09±597.18 Bd | 7 747.71±497.77 Ae | ||||||||
秆下部箨片lower culm blade | 11 901.32±1 911.33 Bb | 8 597.93±742.84 Bc | 5 201.74±535.55 Bd | 5 086.25±912.42 Bd | 18 631.57±3 104.70 Aa | 1 926.95±226.21 Ce | ||||||||
秆上部箨鞘upper culm sheath | 3 528.08±412.34 Dbc | 3 336.13±424.28 Dc | 3 991.63±454.26 Ca | 3 955.65±807.15 Ca | 3 643.39±268.42 Db | 3 270.51±465.34 Bd | ||||||||
秆下部箨鞘lower culm sheath | 6 246.34±539.05 Ca | 6 175.70±923.36 Ca | 3 879.15±455.69 Cc | 2 658.89±276.45 Dd | 5 216.87±619.07 Vb | 1 855.41±112.22 Ce | ||||||||
韧皮部 面积/μm2 phloem area | 叶片leaf blade | 415.90±28.98 Fc | 312.14±37.52 Fe | 381.14±23.34 Gd | 480.44±52.48 Fb | 541.94±88.00 Ea | 460.77±53.93 Eb | |||||||
叶柄petiole | 611.15±100.57 Eb | 270.42±23.70 Fe | 486.26±75.57 Fc | 668.08±105.05 Da | 505.89±89.75 Ec | 447.08±52.94 Ed | ||||||||
叶鞘leaf sheath | 1 922.79±282.44 Ba | 644.52±125.57 Ede | 708.92±165.37 Dd | 587.14±109.84 Ee | 960.51±166.17 Cb | 869.77±121.36 Cc | ||||||||
叶枝twig | 2 536.22±405.44 Aa | 818.33±123.08 Dcd | 866.18±261.10 Dbc | 956.47±123.74 Bb | 916.86±111.17 Cbc | 719.72±94.20 Dd | ||||||||
秆上部箨片upper culm blade | 632.48±142.69 Eb | 628.29±193.31 Eb | 589.41±91.10 Eb | 653.82±161.18 Db | 746.13±157.95 Da | 422.79±87.65 Ec | ||||||||
秆下部箨片lower culm blade | 1 300.45±359.59 Db | 902.64±177.46 Cc | 835.23±132.52 Cc | 577.31±129.68 Ed | 1 977.39±371.37 Aa | 804.71±158.22 Cc | ||||||||
秆上部箨鞘upper culm sheath | 374.14±50.84 Gd | 2 078.86±144.45 Aa | 1 561.56±212.29 Bb | 1 021.82±121.52 Ac | 1 518.81±193.86 Bb | 1 521.85±215.68 Ab | ||||||||
秆下部箨鞘lower culm sheath | 1 560.15±341.33 Cb | 1 194.62±320.41 Bc | 1 906.48±219.37 Aa | 862.79±123.49 Cd | 2 025.80±288.92 Aa | 1 074.85±214.94 Bc | ||||||||
木质部占维管束面积比/% area ratio of xylem to vascular bundle | 叶片leaf blade | 15.51±1.54 Eb | 11.25±2.08 Ed | 9.90±0.92 Fe | 12.30±4.50 Ec | 19.03±3.81 Da | 16.05±2.66 Eb | |||||||
叶柄petiole | 8.85±2.12 Gd | 8.43±2.01 Fd | 10.83±1.76 EFc | 13.72±3.26 Ea | 12.71±2.72 Fb | 14.15±3.22 Fa | ||||||||
叶鞘leaf sheath | 22.21±3.43 Ca | 15.07±3.15 Cc | 16.74±3.77 Cb | 11.08±4.23 Ee | 12.79±1.92 Fe | 15.15±3.55 EFc | ||||||||
叶枝twig | 10.81±4.76 Fd | 14.39±4.25 CDc | 11.31±5.16 Ed | 21.98±5.23 Da | 17.37±2.91 Eb | 11.42±2.32 Gd | ||||||||
秆上部箨片upper culm blade | 40.02±4.25 Aa | 31.56±1.68 Bb | 22.93±1.15 Ac | 29.80±5.79 Cb | 23.69±1.28 Cc | 45.10±4.68 Aa | ||||||||
秆下部箨片lower culm blade | 29.37±4.13 Bc | 42.73±4.07 Aa | 23.62±2.66 Ad | 34.11±5.49 Bb | 44.92±5.33 Aa | 23.75±3.33 Cd | ||||||||
秆上部箨鞘upper culm sheath | 28.75±5.61 Bbc | 13.13±5.27 Dd | 13.18±3.61 Dd | 42.27±6.08 Aa | 26.21±4.11 Bc | 30.08±4.59 Bb | ||||||||
秆下部箨鞘lower culm sheath | 20.77±4.04 Dc | 32.19±3.98 Ba | 19.32±3.62 Bd | 23.03±3.89 Db | 20.34±5.10 Dc | 21.26±3.73 Dbc | ||||||||
韧皮部占维管 束面积比/% area ratio of phloem to vascular bundle | 叶片leaf blade | 16.03±2.21 Cd | 12.70±2.36 BCe | 18.04±2.46 Bb | 17.44±3.84 Cc | 20.12±3.26 Aa | 16.84±3.40 Bcd | |||||||
叶柄petiole | 21.82±3.85 Ba | 13.73±3.19 Bc | 21.98±4.55 Aa | 21.76±4.09 Ba | 17.60±3.67 BCb | 18.64±4.22 Ab | ||||||||
叶鞘leaf sheath | 24.64±3.90 Aa | 16.20±2.42 Ac | 21.26±3.42 Ab | 16.55±6.87 Cc | 16.57±3.90 Cc | 16.16±3.75 Bc | ||||||||
叶枝twig | 14.85±3.13 Dd | 16.10±2.14 Ac | 15.90±4.06 Cc | 26.63±6.07 Aa | 18.75±3.09 Bb | 12.96±2.76 De | ||||||||
秆上部箨片upper culm blade | 5.09±2.20 Fc | 4.16±1.29 Ed | 2.86±0.73 Ge | 8.77±4.76 Ea | 6.65±2.57 Eb | 5.59±1.33 Fbc | ||||||||
秆下部箨片lower culm blade | 3.21±1.40 Gd | 4.92±1.82 Eb | 3.71±1.05 Fc | 3.85±0.90 Fc | 4.84±1.09 Fb | 9.95±2.57 Ea | ||||||||
秆上部箨鞘upper culm sheath | 3.18±0.69 Gd | 11.10±0.73 Cb | 5.66±0.90 Ec | 14.34±3.69 Da | 11.61±3.11 Db | 14.52±3.24 Ca | ||||||||
秆下部箨鞘lower culm sheath | 8.42±2.45 Ec | 6.32±1.76 Dd | 10.36±2.08 Db | 7.54±1.52 Ecd | 6.79±1.83 Ed | 17.27±3.37 ABa | ||||||||
泡状细胞复合 体面积/μm2 motor cell area | 叶片leaf blade | 888.48±86.51 Ca | 798.55±148.48 Cc | 817.92±114.21 Cbc | 820.67±82.30 Cbc | 837.01±98.84 Bab | 855.11±91.12 Bab | |||||||
秆上部箨片upper culm blade | 4 628.46±277.03 Aa | 3 431.74±576.76 Bb | 1 158.58±151.60 Be | 1 247.57±143.08 Bde | 2 591.32±377.24 Ac | 1 367.64±132.83 Ad | ||||||||
秆下部箨片lower culm blade | 1 039.01±169.02 Bd | 4 686.63±642.83 Ab | 5 086.44±1 938.50 Aa | 3 773.65±791.72 Ac | — | — |
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