药用植物博落回形态与发育解剖学研究

doi:10.3724/SP.J.1259.2015.00072

植物学报 ›› 2015, Vol. 50 ›› Issue (1): 72-82.DOI: 10.3724/SP.J.1259.2015.00072

药用植物博落回形态与发育解剖学研究

程巧^1,², 乐捷^2,^*(), 曾建国^1,^*()

¹湖南农业大学, 国家中药材生产(湖南)技术中心, 长沙 410128
²中国科学院植物研究所, 植物分子生理学重点实验室, 北京 100093

收稿日期:2014-02-17 接受日期:2014-04-03 出版日期:2015-01-01 发布日期:2015-04-09
通讯作者: 乐捷,曾建国
作者简介:
? 共同第一作者
基金资助:
国家自然科学基金(No.31200615)、湖南省自然科学基金(No.12JJ4027)和作物种质创新与资源利用国家重点实验室培育基地开放项目(No.11KFXM04)

Morphology and Developmental Anatomy of Medicinal Plant Macleaya cordata

Qiao Cheng^{1, 2}, Jie Le^{2, *}, Jianguo Zeng^{1, *}

¹National Chinese Medicinal Herbs (Hunan) Technology Center, Hunan Agricultural University, Changsha 410128, China
²Key Laboratory of Plant Molecular Physiology, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China

Received:2014-02-17 Accepted:2014-04-03 Online:2015-01-01 Published:2015-04-09
Contact: Le Jie,Zeng Jianguo
About author:
? These authors contributed equally to this paper

摘要/Abstract

摘要： 博落回(Macleaya cordata)是罂粟科含有异喹啉生物碱的主要药用植物, 其生物碱提取物具有多种药理活性。目前对博落回的研究主要集中在生物碱的成分、活性和药理等方面, 对与其合成、运输、储藏密切相关的发育解剖结构及生物碱组织化学定位研究还未见报道。该文报道了温室中生长的博落回植株形态特征, 并结合徒手切片和改良碘化铋钾沉淀的方法对主要器官(根、茎、叶)的发育结构和生物碱组织化学染色进行了初步分析。结果表明, 博落回发育基本遵循一般双子叶植物的规律。根中生物碱出现在中柱鞘外的薄壁细胞和导管周围。茎和叶柄中的生物碱主要出现在维管束周围, 偶尔也出现在茎的髓细胞中。总体上, 茎中的生物碱积累量少于根。

Abstract: Macleaya cordata, in the Papaveraceae family, is a common medicinal plant used for benzylisoquinoline alkaloids and other alkaloids. Current research into M. cordata has focused on the composition, bioactivity and pharmacology of alkaloids. The anatomical structure of M. cordata organs and histochemical localization of alkaloids, closely related to alkaloid synthesis, transport and storage, have not been reported. Here we show the morphologic characters of greenhouse-grown M. cordata plants and developmental anatomic structure of main vegetative organs by hand sectioning and the histochemical localization of alkaloids in roots, stems and leaves by Dragendorff reagent. M. cordata is a typical dicotyledonous plant. Abundant alkaloid accumulation occurred in parenchymal cells and cells around the vessels of stele of roots, cells near the vascular bundles of stems and petioles, and occasionally in the pith of stems. Overall, the amount of alkaloids in roots is higher than that in stems.

程巧, 乐捷, 曾建国. 药用植物博落回形态与发育解剖学研究. 植物学报, 2015, 50(1): 72-82.

Qiao Cheng, Jie Le, Jianguo Zeng. Morphology and Developmental Anatomy of Medicinal Plant Macleaya cordata. Chinese Bulletin of Botany, 2015, 50(1): 72-82.

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图/表 5

图1 博落回植株外部形态与叶片表皮观察 (A)-(D) 博落回不同时期植株形态, 分别为1周(A)、8周(B)、12周(C)和16周(D)植株; (E) 花序模式图; (F) 花苞时期, 示花序局部; (G) 花成熟期, 示花序局部; (H) 花; (I) 雄蕊; (J) 雌蕊; (K) 成熟果荚; (L) 果荚外部形态; (M) 果荚内部形态; (N) 种子外部形态; (O) 种子内部形态; (P) 8周植株第1至第7叶的正面; (Q) 8周植株第1至第7叶的反面; (R) 叶上表皮; (S) 叶下表皮。(R), (S) Bar=100 μm

Figure 1 External morphology and leaf epidermis observation of Macleaya cordata (A)-(D) The overall view of one (A), eight (B), twelve (C)- and sixteen (D)-week old plants; (E) Illustration of inflorescence; (F) Part of young inflorescence; (G) Part of mature inflorescence; (H) Flower; (I) Stamens; (J) Pistil; (K) Mature capsule; (L) External morphology of capsule; (M) Internal morphology of capsule; (N) External morphology of seeds; (O) Internal morphology of seeds; (P) Upper surface of leaf one to seven from eight-week-old seedlings; (Q) Lower surface of leaf from one to seven eight-week-old seedlings; (R) Upper epidermis; (S) Lower epidermis. (R), (S) Bar=100 μm

图2 博落回根的发育解剖结构 (A) 博落回植株幼苗, 示根横切取样点(箭头所示); (B) 4周根的初生结构, 示表皮、皮层和中柱; (C) 6周根的初生结构, 示初生到次生结构的过渡; (D) 8周根的次生结构, 示后生木质部导管; (E) 10周根的次生结构, 示维管射线; (F) 12周根的次生结构, 示维管形成层; (G) 13周根的次生结构, 示木纤维; (H) 14周根的次生结构, 示次生木质部和次生韧皮部; (I) 16周根的次生结构, 示中柱鞘; (J) 20周根的次生结构, 示周皮裂缝(箭头所示)。切片中橘黄色斑点疑为生物碱积累(红色箭头所示)。e: 表皮; c: 皮层; st: 中柱; pxy: 初生木质部; pph: 初生韧皮部; mv: 后生木质部导管; vr: 维管射线; vca: 维管形成层; xf: 木纤维; sxy: 次生木质部; sph: 次生韧皮部; pe: 中柱鞘环。(B)-(H) Bar=100 μm; (I) Bar=250 μm; (J) Bar=300 μm

Figure 2 Developmental anatomy of Macleaya cordata roots (A) Seedlings of Macleaya cordata, an arrow points to the position of section; (B) The cross section shows the primary structure of a 4-week-old root, shows the epidermis, cortex and stele; (C) The cross section of a 6-week-old root shows the primary structure to secondary structure; (D) The secondary structure of a 8-week-old root shows the mature metaxylem vessels; (E) The secondary structure of a 10-week-old root shows vascular ray; (F) The secondary structure of a 12-week-old root shows vascular cambium; (G) The secondary structure of a 13-week-old root, showing xylon fiber; (H) The secondary structure of 14-week-old root shows secondary xylem and secondary phloem; (I) The secondary structure of a 16-week-old root shows pericycle; (J) The secondary structure of a 20-week-old root, arrows indicate the crevices in periderm (arrows); Oranges dots in sections are putative accumulation of alkaloids (red arrows). e: Epidermis; c: Cortex; st: Stele; pxy: Primary xylem; pph: Primary phloem; mv: Metaxylem vessels; vr: Vascular ray; vca: Vascular cambium; xf: Xylon fiber; sxy: Secondary xylem; sph: Secondary phloem; pe: Pericycle. (B)-(H) Bar=100 μm; (I) Bar=250 μm; (J) Bar=300 μm

图3 博落回茎的发育解剖结构 (A) 博落回植株的一部分, 示茎取样点(箭头所示); (B) 6周茎的初生结构, 示表皮、皮层、髓和维管束; (C) 茎表皮; (D) 茎的局部, 示表皮和皮层; (E) 8周茎的初生结构, 示原生木质部导管和原生韧皮部筛管的分化; (F) 12周茎, 示初生木质部、初生韧皮部和维管形成层; (G) 16周茎的次生结构, 示韧皮纤维; (H) 20周茎, 示维管射线。切片中橘黄色斑点疑为生物碱积累(红色箭头所示)。Pi: 髓; pr: 髓射线; e: 表皮; c: 皮层; vab: 维管束; v: 木质部导管; st: 韧皮部筛管; vca: 维管形成层;; pxy: 初生木质部; pph: 初生韧皮部; pf: 韧皮纤维。(B)-(D) Bar=100 μm; (E)-(H) Bar=200 μm

Figure 3 Developmental anatomy of Macleaya cordata stem (A) Part of Macleaya cordata seedlings, an arrow points to the position of section; (B) The cross section of a 6-week-old primary meristem in the stem shows the cortex, epidermis, pith and vascular bundle; (C) Epidermis of stem; (D) Enlarged image shows the epidermis and cortex; (E) The cross section of a 8-week-old primary meristem in the stem shows the differentiation of protoxylem vesseIs and protophloem sieve tubes; (F) The cross section of a 12-week-old stem shows primary xylem, primary phloem and vascular cambium of stem; (G) Secondary structure of a 16-week-old stem shows phloem fiber; (H) The structure of a 20-week-old stem shows vascular ray. Oranges dots in sections are putative accumulation of alkaloids (red arrows). Pi: Pith; pr: Pith ray; e: Epidermis; c: Cortex; vab: Vascular bundle; v: VesseIs; st: Sieve tubes; vca: Vascular cambium; pxy: Primary xylem; pph: Primary phloem; pf: Phloem fiber. (B)-(D) Bar=100 μm; (E)-(H) Bar=200 μm

图4 博落回叶片和不同叶位叶柄的结构 (A) 切片位点, 蓝色箭头所示为叶片处横切(图B, C), 叶柄横切处如橘色箭头从上至下所示, 叶柄靠近叶片(图D, G, J)、叶柄中部(图E, H, K)和叶柄靠近茎(图F, I, L); (B) 异面叶, 示已分化出栅栏组织和海绵组织; (C) 叶脉, 示维管束中的维管形成层、木质部、韧皮部和韧皮纤维; (D)-(F) 分别示第1叶位叶柄不同位置的横切面; (G)-(I) 分别示第5叶位叶柄不同位置的横切面; (J)-(L) 分别示第10叶位叶柄不同位置的横切面; 切片中橘黄色细胞疑为生物碱积累所致, 如红色箭头所示。e: 表皮; c: 皮层; vab: 维管束; vca: 维管形成层; pf: 韧皮纤维; xy: 木质部; ph: 韧皮部; eh: 表皮毛; pa: 栅栏组织; s: 海绵组织。(B)-(I) Bar=200 μm; (J) Bar=50 μm; (K) Bar=100 μm; (L) Bar=1 mm

Figure 4 The structure of Macleaya cordata leaf and petiole (A) Image of a leaf; Arrows from bottom to the top point to the positions of section at petioles near blade (panel D, G, J), middle (panel E, H, K), near stem (panel F, I, L) and leaf blade (panel B and C); (B) Palisade and spongy cells in leaves; (C) Vascular bundles include vascular cambium, xylogen, phloem and phloem fiber; (D)-(F) The cross sections of different location on the first petioles; (G)-(I) The cross section of different location on the fifth petioles; (J)-(L) The cross section of different location on the tenth petioles; Orange coloured cells in sections are putative accumulation of alkaloids (red arrows). e: Epidermis; c: Cortex; vab: Vascular bundle; vca: Vascular cambium; pf: Phloem fiber; xy: Xylogen; ph: Phloem; eh: Epidermal hair; pa: Palisade; s: Spongy. (B)-(I) Bar=200 μm; (J) Bar=50 μm; (K) Bar=100 μm; (L) Bar=1 mm

图5 博落回根、茎和叶柄生物碱的组织定位 (A)-(C) 分别为6、12和20周根未经染色切片照片; (D)-(F) 分别为以上切片经改良碘化铋钾染色后的照片; (G)-(I) 分别为6、12和20周茎未经染色切片照片; (J)-(L) 分别为以上切片经改良碘化铋钾染色后的照片; (M)-(O) 分别为靠近叶片处、中部及靠近茎处叶柄横切片, 经改良碘化铋钾染色后照片; (P)-(R) 紫外激发荧光法验证生物碱的组织定位, 根的横切片明场照片(P); 紫外激发荧光照片(Q); 改良碘化铋钾处理后各种紫外激发荧光明显减弱(R); (S)-(U) 生物碱溶剂处理对生物碱紫外荧光的影响, 根横切片明场(S); 紫外激发荧光(T); 溶剂处理后紫外激发荧光消失, 但细胞壁自发荧光仍存在(U)。照片中箭头指示生物碱富集区。 (A) Bar=100 μm; (B), (C) Bar=200 μm; (D) Bar=100 μm; (E), (F) Bar=200 μm; (G)-(L) Bar=100 μm; (M)-(O) Bar=200 μm; (P)-(U) Bar=100 μm

Figure 5 Histochemical localization of alkaloids in Macleaya cordata roots, stems and petioles (A)-(C) Cross sections from 6-, 12-, and 20-week-old roots, respectively; (D)-(F) Images of above root samples stained with Dragendorff reagents; (G)-(I) Cross sections from the 6-, 12-, and 20-week-old stems, respectively; (J)-(L) Images of above stem samples stained with Dragendorff reagents; (M)-(O) The stained sections of a petiole at the position near blade, middle, and near stem; (P)-(R) Alkaloide histochemical localization confirmed by UV-fluorescence; A bright-field image of a root section (P), Fluorescent image after UV stimulation (Q); UV-stimulated fluorescence were not observed after Dragendorff reagent staining (R); (S)-(U) Solvent treatment abolished the UV-stimulated fluorescence; A bright-field image of a root section (S), Image of UV-stimulated fluorescence (T). Fluorescence from alkaloid accumulations were abolished by a treatment with a mixed solvent of 1% hydrochloric acid: methyl alcohol (U). Blue arrows point to the alkaloid accumulations. (A) Bar=100 μm; (B), (C) Bar=200 μm; (D) Bar=100 μm; (E), (F) Bar=200 μm; (G)-(L) Bar=100 μm; (M)-(O) Bar=200 μm; (P)-(U) Bar=100 μm

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药用植物博落回形态与发育解剖学研究

Morphology and Developmental Anatomy of Medicinal Plant Macleaya cordata

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