药学学报

Latin name	Height /cm	Adaxial side	Abaxial side
Anoectochilus roxburghii	8-18	Dark green to blackish purple	Ovate to ovate-orbicular, 1.3-4.5 × 0.8-3.6 cm, apex acute	Pinkish golden reticulate venation	Pale purplish red
Anoectochilus formosanus	11-20	Dark green	Ovate to ovate-orbicular, 2-5 × 1-3.5 cm, apex acute	White venation	Reddish purple
Anoectochilus chapaensis	12-18	Blackish green	Ovate, oblique, 2-5 × 1.5-3.5 cm, apex acute	Golden-red venation	Pale green
Anoectochilus koshunensis	about 20	Dark green	Ovate to ovate-orbicular, 3-4 × 2-3 cm	White venation	Reddish purple
Anoectochilus lanceolatus	15-30	Dark green	Ovate, ovate-lanceolate, or elliptic, 1.5-8 × 1-5 cm, apex acute	Whitish stripes along midvein and 2 lateral veins	Pale green
Anoectochilus elwesii	15-25	Dark purple or blackish green	Ovate to ovate-orbicular, 1.5-5 × 1-3 cm	3 red veins sometimes	Pale red or pale green
Anoectochilus moulmeinensis	20-30	Green	Oblong to narrowly elliptic, 5-8 × 2-3 cm	Whitish and broad stripes along midvein	Greyish green
Anoectochilus zhejiangensis	8-16	Dark greenish purple	Broadly ovate to ovate-orbicular, 0.7-2.6 × 0.6-2.1 cm, margin slightly undulate, apex acute	Golden-red venation	Pale purplish red
Anoectochilus pingbianensis	15-18	Dark green	Ovate, 3-4.5 × 2-3 cm	Pinkish golden reticulate venation	Pale red or pale green
Anoectochilus burmannicus	16-30	Blackish green	Broadly ovate to ovate-elliptic, slightly oblique, 3-8 × 1.8-5 cm, apex acute	Limited bronze venation	Pinkish red
Anoectochilus emeiensis	19-21	Blackish green	Ovate, 3.5-4 × 3-3.2 cm	Pinkish golden reticulate venation	Reddish purple
Anoectochilus yungianus	10-20	Dark green	Ovate, 1.5-5 × 1-3 cm	Pinkish golden reticulate venation	Dark purple

Latin name	Height /cm	Adaxial side	Abaxial side
Anoectochilus roxburghii	8-18	Dark green to blackish purple	Ovate to ovate-orbicular, 1.3-4.5 × 0.8-3.6 cm, apex acute	Pinkish golden reticulate venation	Pale purplish red
Anoectochilus formosanus	11-20	Dark green	Ovate to ovate-orbicular, 2-5 × 1-3.5 cm, apex acute	White venation	Reddish purple
Anoectochilus chapaensis	12-18	Blackish green	Ovate, oblique, 2-5 × 1.5-3.5 cm, apex acute	Golden-red venation	Pale green
Anoectochilus koshunensis	about 20	Dark green	Ovate to ovate-orbicular, 3-4 × 2-3 cm	White venation	Reddish purple
Anoectochilus lanceolatus	15-30	Dark green	Ovate, ovate-lanceolate, or elliptic, 1.5-8 × 1-5 cm, apex acute	Whitish stripes along midvein and 2 lateral veins	Pale green
Anoectochilus elwesii	15-25	Dark purple or blackish green	Ovate to ovate-orbicular, 1.5-5 × 1-3 cm	3 red veins sometimes	Pale red or pale green
Anoectochilus moulmeinensis	20-30	Green	Oblong to narrowly elliptic, 5-8 × 2-3 cm	Whitish and broad stripes along midvein	Greyish green
Anoectochilus zhejiangensis	8-16	Dark greenish purple	Broadly ovate to ovate-orbicular, 0.7-2.6 × 0.6-2.1 cm, margin slightly undulate, apex acute	Golden-red venation	Pale purplish red
Anoectochilus pingbianensis	15-18	Dark green	Ovate, 3-4.5 × 2-3 cm	Pinkish golden reticulate venation	Pale red or pale green
Anoectochilus burmannicus	16-30	Blackish green	Broadly ovate to ovate-elliptic, slightly oblique, 3-8 × 1.8-5 cm, apex acute	Limited bronze venation	Pinkish red
Anoectochilus emeiensis	19-21	Blackish green	Ovate, 3.5-4 × 3-3.2 cm	Pinkish golden reticulate venation	Reddish purple
Anoectochilus yungianus	10-20	Dark green	Ovate, 1.5-5 × 1-3 cm	Pinkish golden reticulate venation	Dark purple

Latin name	Color	Sepal	Lip resupinate	Lip	Calli	Florescence
A. roxburghii	Pinkish white	Pinkish, outer surface pubescent, 1-veined	No	Pinkish white, Y-shaped; epichile 2-lobed, lobes diverging at an acute angle to one another, obovate-oblong to spatulate; spur conic, apex shallowly bilobed	2 irregular calli, fleshy near entrance of spur	August-December
A. formosanus	White	Reddish brown, pubescent on outer surface	Yes	Pale yellowish white, Y-shaped; epichile longitudinally dilated; spur conic, apex shallowly bilobed	2 calli, near the end of spur, plate	October-November
A. chapaensis	White	Pubescent on outer surface, 1-veined	No	White, Y-shaped; epichile 2-lobed, lobes diverging widely, obovate, margin crenulate, apex truncate; spur conic, apex obtuse	2 calli located near the middle of spur	July-August
A. koshunensis	White	Reddish brown, pubescent on outer surface	No	White, Y-shaped; epichile 2-lobed, slightly brown, margin entire or slightly crenulate; spur conic, extending upward beyond the lobes	2 calli, fleshy near entrance of spur	July-October
A. lanceolatus	Yellow	Yellowish green, glabrous, 1-veined	Yes	Golden yellow, Y-shaped; epichile 2-lobed, cuneate-oblong to obovate, margin entire or slightly crenulate, apex truncate	Slender, needle-like, linear, lanceolate, often hooked upward, sessile	June-September
A. elwesii	White	Green or white	Yes	White, Y-shaped; epichile 2-lobed; nearly obovate; margin entire or slightly crenulate, glabrous	A seperate longitudinal fold, concave near square at top	July-August
A. moulmeinensis	Pinkish red	Pinkish red, outer surface pubescent, 1-veined	Yes	T-shaped, margin slightly crenulate; epichile 2-lobed, nearly obovate; short, margin entire	A longitudinal fold; 2 ovoid, short stipe, apically serrated	August-October
A. zhejiangensis	White	Pinkish red, outer surface pubescent	No	White, Y-shaped; epichile 2-lobed, lobes diverging at an acute angle to one another, flabellate, slightly obliquespur conic, slightly curved, apex shallowly bilobed	2 irregular, stalked calli near entrance	July-September
A. pingbianensis	White	Pinkish	Yes	White, T-shaped; extending forward; epichile 2-lobed, inverted triangle; spur conic	2 calli, fleshly, narrowly elliptic, with short stalk	October
A. burmannicus	Yellowish white	Pink, outer surface pubescent, 1-veined	No	Yellow, Y-shaped; epichile 2-lobed, lobes diverging at an acute angle to one another, oblong, oblanceolate, or spatulatespur conic, apex obtuse	2 subelliptic, sessile calli near entrance	September-December
A. emeiensis	White with purplish red	Fuchsia	No	White, Y-shaped, apex shallowly bilobe; epichile 2-lobed, oblong, oblanceolate; spur straight	2 calli, nearly cuneate	August-September
A. yungianus	White	Greyish green with pinkish, outer surface pubescent	No	White, Y-shaped; epichile 2-lobed, inverted triangle; spur conic	2 calli, near the middle of spur	February

Latin name	Color	Sepal	Lip resupinate	Lip	Calli	Florescence
A. roxburghii	Pinkish white	Pinkish, outer surface pubescent, 1-veined	No	Pinkish white, Y-shaped; epichile 2-lobed, lobes diverging at an acute angle to one another, obovate-oblong to spatulate; spur conic, apex shallowly bilobed	2 irregular calli, fleshy near entrance of spur	August-December
A. formosanus	White	Reddish brown, pubescent on outer surface	Yes	Pale yellowish white, Y-shaped; epichile longitudinally dilated; spur conic, apex shallowly bilobed	2 calli, near the end of spur, plate	October-November
A. chapaensis	White	Pubescent on outer surface, 1-veined	No	White, Y-shaped; epichile 2-lobed, lobes diverging widely, obovate, margin crenulate, apex truncate; spur conic, apex obtuse	2 calli located near the middle of spur	July-August
A. koshunensis	White	Reddish brown, pubescent on outer surface	No	White, Y-shaped; epichile 2-lobed, slightly brown, margin entire or slightly crenulate; spur conic, extending upward beyond the lobes	2 calli, fleshy near entrance of spur	July-October
A. lanceolatus	Yellow	Yellowish green, glabrous, 1-veined	Yes	Golden yellow, Y-shaped; epichile 2-lobed, cuneate-oblong to obovate, margin entire or slightly crenulate, apex truncate	Slender, needle-like, linear, lanceolate, often hooked upward, sessile	June-September
A. elwesii	White	Green or white	Yes	White, Y-shaped; epichile 2-lobed; nearly obovate; margin entire or slightly crenulate, glabrous	A seperate longitudinal fold, concave near square at top	July-August
A. moulmeinensis	Pinkish red	Pinkish red, outer surface pubescent, 1-veined	Yes	T-shaped, margin slightly crenulate; epichile 2-lobed, nearly obovate; short, margin entire	A longitudinal fold; 2 ovoid, short stipe, apically serrated	August-October
A. zhejiangensis	White	Pinkish red, outer surface pubescent	No	White, Y-shaped; epichile 2-lobed, lobes diverging at an acute angle to one another, flabellate, slightly obliquespur conic, slightly curved, apex shallowly bilobed	2 irregular, stalked calli near entrance	July-September
A. pingbianensis	White	Pinkish	Yes	White, T-shaped; extending forward; epichile 2-lobed, inverted triangle; spur conic	2 calli, fleshly, narrowly elliptic, with short stalk	October
A. burmannicus	Yellowish white	Pink, outer surface pubescent, 1-veined	No	Yellow, Y-shaped; epichile 2-lobed, lobes diverging at an acute angle to one another, oblong, oblanceolate, or spatulatespur conic, apex obtuse	2 subelliptic, sessile calli near entrance	September-December
A. emeiensis	White with purplish red	Fuchsia	No	White, Y-shaped, apex shallowly bilobe; epichile 2-lobed, oblong, oblanceolate; spur straight	2 calli, nearly cuneate	August-September
A. yungianus	White	Greyish green with pinkish, outer surface pubescent	No	White, Y-shaped; epichile 2-lobed, inverted triangle; spur conic	2 calli, near the middle of spur	February

我国金线兰资源特征及繁育技术研究进展

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吕欣锴 , 周丽思 ^* , 郭顺星 ^*

药学学报 | 综述 2022,57(7): 2057-2067

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药学学报 | 综述 2022, 57(7): 2057-2067

我国金线兰资源特征及繁育技术研究进展

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吕欣锴, 周丽思^*, 郭顺星^*

作者信息

中国医学科学院、北京协和医学院药用植物研究所, 北京 100193

通讯作者:

*周丽思, E-mail: zls921203@sina.com;

郭顺星, E-mail: sxguo2015@163.com

Resource characteristics and propagation techniques of Anoectochilus roxburghii in China

Xin-kai LÜ, Li-si ZHOU^*, Shun-xing GUO^*

Affiliations

Institute of Medicinal Plant Development, Chinese Academy of Medical Science and Peking Union Medical College, Beijing 100193, China

出版时间: 2022-07-12 doi: 10.16438/j.0513-4870.2022-0019

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摘要

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金线兰(Anoectochilus roxburghii) 是兰科开唇兰属药用物种, 其全草的鲜品或干品可作为“金线莲”药材使用, 具有降血糖、降血脂、抗炎镇痛等功效。在广东、福建等地区的食药用历史悠久, 延续至今。随着“药食同源”概念的广泛认同、市场需求激增, 野生金线兰资源已远不能满足供应, 处于濒危状态, 市场价格逐年推高。建立人工繁育体系, 是缓解野生资源紧缺的重要手段。明确金线兰资源特征, 则是优选种质资源、优化繁育体系、建立有效评价标准的关键依据。因此, 本文从本草考证、资源分布、资源鉴定、化学成分特征与药理活性、繁育技术进行综述, 旨在为我国金线兰资源可持续利用和后续深入研究提供参考。

关键词

金线兰 / 资源特征 / 繁育技术 / 菌根共生 / 兰科

Abstract

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Anoectochilus roxburghii (Wall.) Lindl. is a medicinal species belonging to the Orchidaceae, whose whole plant can be used as a medicinal herb, known as "JinXianLian". It has antidiabetic, liver-protecting, anti-inflammatory, etc. A. roxburghii has long been used as food and medicine in Guangdong and Fujian provinces. With the wide recognition of the concept of "medicine and food homology" and the surge of market demand, wild A. roxburghii has been far from meeting the supply. It is important to establish an artificial propagation system. Resource characteristics are the key basis for optimizing germplasm and propagation systems. Therefore, this paper summarizes the germplasm resource characteristics and propagation technologies of A. roxburghii in China to provide a reference for sustainable development and subsequent mechanistic research.

Key words

Anoectochilus roxburghii / resource characteristics / propagation technique / mycorrhizal symbiosis / Orchidaceae

引用本文

吕欣锴, 周丽思, 郭顺星. 我国金线兰资源特征及繁育技术研究进展. 药学学报, 2022 , 57 (7) : 2057 -2067 . DOI: 10.16438/j.0513-4870.2022-0019

Xin-kai LÜ, Li-si ZHOU, Shun-xing GUO. Resource characteristics and propagation techniques of Anoectochilus roxburghii in China[J]. Acta Pharmaceutica Sinica, 2022 , 57 (7) : 2057 -2067 . DOI: 10.16438/j.0513-4870.2022-0019

正文

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金线兰[Anoectochilus roxburghii (Wall.) Lindl.], 又名花叶开唇兰, 是兰科(Orchidaceae) 开唇兰属(Anoectochilus) 地生型植物, 药材“金线莲”的基源植物之一。根据《中国植物志》 (2004年版) 的统计, 全球开唇兰属植物共有40余种, 其中我国存在20个种及2个变种。此后, 陆续有新种在我国被首次发现, 如麻栗坡金线兰、丽蕾金线兰等^[1-4]。金线兰民间应用历史悠久, 其全草入药, 具有降血糖、清热凉血、除湿解毒、保肝、抗炎镇痛等功效。主要分布于福建、浙江、广东、云南等南部省份。野生金线兰资源较为稀缺, 《濒危野生动植物种国际贸易公约》 (CITES) 将其列入附录Ⅱ的保护物种, 《国家重点保护野生植物名录》 (第二批) 将其列为二级保护植物。随着人工繁育技术的发展, 金线兰产业得到了迅速地发展, 而明确金线兰资源特征和繁育特性将直接影响该产业的生命力。因此, 本文对金线兰资源特征和可持续利用的相关研究进行综述, 旨在为金线兰资源理性开发及可持续发展提供指导。

1 本草考证

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金线兰于1839年被J.F. Royle首次记载, 在我国记载时间较晚。古代经典药用植物著作如《神农本草经》、《本草纲目》等中均无记载。目前可追溯的最早记载是出版于1976年的《中国高等植物图鉴》, 并首次提到其具有药用价值, 也进一步证实金线兰自古便资源稀少, 生长于人迹罕至的深山中。金线兰别名众多, 叫法混杂。据《全国中草药名鉴》中记载, “金石松”、“金蚕”、“金石蚕”、“少年红”、“树草莲”、“鸟人参”、“金线虎头蕉”、“金线入骨消”等均为金线兰的别名。此外, 不同地区对金线兰叫法也有差异, 如海南省称之为“金钱枫”; 在四川省峨眉山地区叫“蛇皮兰”。甚至同一省份不同地区也有不同称呼, 如在广西贺县称之为“小叶金耳环”, 在桂平县被叫做“金丝线”, 而在鹿寨县又叫“麻叶菜”。需要注意的是, 文献和著作中常将台湾银线兰(Anoectochilus formosanus Hayata.) 也称为“金线兰”, 造成这种现象的原因: ①台湾省“金线兰”官方所指植物与其他省份所指非同种, 台湾省的“金线兰”特指台湾银线兰, 而根据《中国植物志》所载, 其他省的“金线兰”特指花叶开唇兰。②在《中国植物志》出版之前, 未对“金线兰”进行统一界定, 因此此前出版的著作, 如《中华本草》、《全国中草药名鉴》等均将金线兰和台湾银线兰作为药材“金线莲”的基源植物。目前国内金线莲市场中, 金线兰或台湾银线兰均得到认可, 两者都具有良好的降血糖、降血脂、抗炎等活性^{[5, 6]}。除此之外, 恒春银线兰(Anoectochilus koshunensis Hayata.)、滇越金线兰(Anoectochilus chapaensis Gagnep.)、齿唇兰(Anoectochilus lanceolatus Lindl.)、西南齿唇兰[Anoectochilus elwesii (Clarke ex Hook.f.) King & Pantl.]、艳丽齿唇兰[Anoectochilus moulmeinensis (Par. & Rchb.f.) Seidenf.] 等5种开唇兰属植物也被证实具有一定药用价值^[7], 但是否能够作为金线兰的替代品, 仍待进一步研究。

2 中国野生资源分布

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金线兰具有环境适应性差、自然生长缓慢、天然繁殖率低的特点。在我国分布在福建、浙江、广西、广东、江西、四川、云南、贵州、海南、湖南、西藏等省份, 尤其以东南省份福建、浙江居多。总体而言, 针对金线兰进行资源普查的省份较少, 只有福建、广西、浙江、贵州及广东等对金线兰进行资源调查。但是可以确定各省的金线兰均处于濒危状态, 主要分布在常绿落叶阔叶林、竹林及谷沟阴湿处。

福建省作为金线兰的主产地之一, 全省境内均有金线兰的分布, 同时也有浙江金线兰分布。福建省内野生金线兰存在两种形态, 一种叶片具网状花纹, 而另一种叶片则无网纹。经花器解剖和理化鉴定, 确定两种形态均为金线兰^[8]。民间将叶片具网状金红色叶脉的金线兰称为“金线兰母”, 该形态数量较多; 而叶片无网状叶脉的金线兰被称为“金线兰公”, 数量较少, 二者往往以伴生形式生长。福建野生金线兰主要栖息于海拔350~950 m山地之间的竹林、阔叶林或竹阔混生林, 通常零星分布在枯枝落叶层或沟边冲积处。据2016年调查, 学者通过走访调查和实地考察相结合, 在闽东适合金线兰生长的代表性山区找到了具有野生金线兰生长的28个样地, 平均每2 000 m²样地内野生金线兰数量仅为28株, 并且呈现逐年减少的趋势^[9]。

广西省除西部石灰岩地区外, 大部分常绿阔叶林地区都有金线兰分布。虽分布区域广, 但数量十分有限。主要栖息于高郁闭度、光照强度在350~2 000 lx之间, 空气相对湿度在70%~85%之间的山涧常绿阔叶林的枯枝落叶层和腐殖质层中。乔木覆盖度可达70%~90%, 树种主要包括壳斗科、桑科、樟科、木兰科、五加科植物。除常绿阔叶树林外, 毛竹针叶树混交林及灌丛也有分布^[10]。

浙江省与福建省类似, 自北向南均有金线兰和浙江金线兰分布, 但二者都处于濒危状态, 部分居群小于5株。山阴面常绿阔叶林下分布较多。当林下土壤团粒结构好、具有较高含量的速效磷、速效钾及水解性氮时, 金线兰分布可连成片。但在岩石和苔藓上零星分布的金线兰则往往植株矮小、独立生长、未能成片^[11]。

贵州省已报道分布有金线兰和台湾银线兰, 主要在兴义、安龙、望谟等地的喀斯特山区, 分布海拔在300~1 200 m, 尤以300~600 m区间分布最多^[12]。主要栖息在郁闭度达到80%以上的亚热带次生性常绿阔叶落叶混交林, 湿度在80%~95%之间, 腐殖质土厚度可达10~20 cm, 并具有较好的透水、透气性^[13]。

此外, 广东省肇庆地区也有报道金线兰和台湾银线兰分布^[14]。主要分布于海拔200~800 m的针阔叶混交林下, 其自然居群中已不足15株, 属于当地濒危物种^[15], 且长势与土层的厚度无相关性。

3 资源鉴定

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金线兰暂未列入《中国药典》, 因此缺乏国家统一的质量控制标准与管理规范, 不同地方标准存在较大差异与不足, 造成市场缺乏有效监管, 质量控制把关以及药材来源追溯困难。随着色谱与分子测序等技术的发展, 金线兰与近似种的鉴定手段愈加丰富, 鉴定的准确度和速度均得到较大提升, 如化学成分鉴别、分子标记法等, 此外, 将多种鉴定技术相结合也成为当前鉴定的一大趋势。

3.1 形态特征及显微鉴定技术

市面上普遍存在金线兰与其他种混用的现象, 主要混淆品包括金线兰近属植物, 如血叶兰; 或同属不同种植物, 如浙江金线兰、滇越金线兰等。不同属植物基本可以从茎、叶等外观形态加以区分, 而同属不同种的植物则更难鉴定, 除了叶片形态、颜色、叶脉, 往往还需要利用花形加以区分。形态及显微鉴定技术是指通过肉眼或显微镜观察外部形态或内部细胞及细胞内含物形态的技术。金线兰部分同属不同种植物之间的形态特征见表 1与表 2, 主要鉴别点包括叶片与叶脉的颜色; 花瓣颜色、唇瓣是否倒置、萼片颜色等^[16]。表 1归纳了叶的主要鉴别点, 如根据叶色, 金线兰、浙江金线兰、西南齿唇兰等为较独特的黑紫或绿紫色, 区别于其余种的暗绿色; 根据叶脉颜色可区分金线兰(金红色网纹) 与台湾银线兰、恒春银线兰(白色); 此外, 金线兰的中叶脉下表面呈半圆形凸出, 滇越金线兰与台湾银线兰两种植物的则呈微凸^[17]。表 2总结了花的形态特征。以唇瓣为例, 除屏边金线兰外, 其余几种金线兰唇瓣均未倒置。而利用唇瓣是否伸出于两侧萼片基部之外则可将台湾银线兰、恒春银线兰与齿唇兰、西南齿唇兰、艳丽齿唇兰进行区分; 还可通过萼片颜色区分金线兰(红色) 与西南齿唇兰(绿色或白色)。尽管形态及显微鉴定是最直观、最直接的鉴定技术, 但对于易混淆品, 如金线兰与浙江金线兰, 仍存在鉴定准确性低、鉴定难度大的问题, 且对未开花植株的鉴定存在较大限制。

3.2 化学鉴定技术

化学成分鉴别是指利用化学手段, 基于药材化学成分种类与含量的差异进行鉴定的技术。金线兰中最常用的化学鉴定技术是指纹图谱技术。其不仅可用于金线兰真伪品的鉴定, 甚至可区分不同种质及种植方式的金线兰。通过建立组培、栽培及野生金线兰的化学指纹图谱, 发现以金线莲苷为代表的强极性区峰的数量包括响应各种栽培方式间无显著差异; 而在其他糖苷类出峰的极性区, 栽培和野生金线兰的峰响应高于组培种; 在芦丁、水仙苷等出峰的中极性区, 栽培种峰响应显著高于野生与组培种; 在弱极性区, 栽培品的槲皮素、山柰酚峰响应最高, 组培的最低, 最后通过比较金线兰、台湾银线兰与血叶兰的指纹图谱, 发现金线兰与台湾银线兰在一定程度上可互相替代; 而血叶兰成分存在较大差异, 因此不可作为金线兰的替代品^[18]。此外, 为提高鉴定准确度, 还可采取将多种鉴定技术相结合方式。Gao等^[19]发现利用蛋白质组学及表面增强激光解吸电离-飞行时间质谱(SELDI-TOF-MS) 指纹图谱, 可解决ITS无法鉴定到种的问题, 从而对福建、台湾、西藏、云南采集的野生开唇兰属及斑叶兰属共5个种的植物进行物种鉴定与区分。化学鉴定可在一定程度上弥补显微鉴定的不足, 但由于化学成分随组织发育时间与空间而变化, 因而化学鉴定技术易受时空等环境因素影响。

3.3 分子标记技术

分子标记技术是一种通过检测生物体在基因或基因型上所产生的变异来反映生物个体之间差异的方法, 其最大的特点在于可不受组织发育时间与空间的限制, 不受环境的影响, 稳定性较高。常见的如限制性片段长度多态性技术(restriction fragment length polymorphism, RFLP)、随机扩增多态性DNA标记技术(random amplified polymorphic DNA, RAPD)、简单重复序列间区(inter-simple sequence repeat, ISSR)、扩增片段长度多态性(amplified fragment length polymorphism, AFLP)、单链构象多态性(single strand conformation polymorphism, SSCP) 等。随着分子生物学及测序技术的快速发展, 以及测序成本的降低, 越来越多的分子标记相关技术被用于药用植物的鉴定^{[20, 21]}。分子标记技术不仅可对不同产地和不同培养方式的金线兰的遗传特征进行分析, 从而实现对其分类与鉴定^[22-24], 还可以检测不同产地的野生金线兰及其近似种的多样性^[25]。由于分子标记技术建立于大量样本之上, 且成本较高, 因此, 现有分子标签的区分效果仍有待进一步验证, 相较于农作物, 分子标记技术在金线兰中应用仍处于初级研究阶段。目前, 学者正致力于开发更简便、高效的标记技术^[26], 使分子标记技术更准确地应用于金线兰的种质资源鉴定。

4 化学成分特征及其药理活性

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金线兰中已鉴定出上百种化合物, 主要包括黄酮、黄酮苷、有机酸、甾体、萜类、生物碱等(图 1), 其中60余种已分离得到, 其余均通过定性或定量方法被检测到。不同化学成分决定了金线兰多方面的药理作用, 包括肝脏保护、降血糖、抗炎、抗氧化、镇痛等。但由于野生金线兰资源濒危, 且市场价格极高, 因此, 除少数体外酶活实验外, 几乎没有药理实验采用野生金线兰作为实验材料, 多采用组培或人工种植苗。

4.1 金线莲苷及其类似物

金线莲苷(kinsenoside; C₁₀H₁₆O₈; 3-(R)-3-β-D-glucopyranosyloxybutanolide) 最早于1993年由日本学者从兰科开唇兰属植物恒春银线兰(Anoectochilus koshunensis) 中分离^[27], 是由3-(R)-羟基-γ-丁内酯与β-D-吡喃葡萄糖经过糖苷键连接而成。随着研究的深入, 陆续从其他物种中检测出金线莲苷, 主要包括开唇兰属的金线兰和台湾银线兰等; 其他属的如番红花(Crocus sativus L.)、斑叶兰(Goodyera schlechtendaliana Rchb. F.) 等。除金线莲苷外, 还从金线兰中分离得到部分金线莲苷类似物(结构见图 1A), 包括4-β-D-吡喃葡萄糖氧基-丁酸甲酯^[28]、kinsenbenol^[29]、kinsendioside A、kinsendioside B^[30]、(S)-3-O-对羟基苄基-γ-丁内酯、(S)-3-羟基-γ-丁内酯^[31]、金线兰素A^[32]。作为金线兰中含量最高的活性成分, 金线莲苷具有显著的降血糖、肝脏保护活性。研究表明, 金线莲苷可以提高链脲佐菌素致糖尿病大鼠的糖耐量, 修复胰岛损伤细胞并改善其功能, 从而发挥治疗糖尿病的功效^[33]。金线莲苷还具有PTP1B抑制活性, 有望作为潜在的PTP1B抑制剂, 用于治疗2型糖尿病^[34]。Xiang等^[35]通过半刀豆球蛋白A诱导小鼠T细胞介导的肝炎模型, 发现金线莲苷可以显著改善自发免疫性肝炎, 降低炎症细胞浸润和促炎因子的生成。通过靶向VEGFR2诱导对自身免疫性肝损伤的免疫抑制, 减少代谢相关PI3K-AKT和炎症相关JAK2-STAT3通路相互作用, 从而破坏树突细胞诱导的CD8⁺T细胞反应的交叉启动。此外, 金线莲苷还可以有效保护视网膜色素上皮细胞免受过氧化氢诱导的氧化应激引起的调亡, 并通过抑制p65、Erk和p38的表达, 减少凋亡相关血管生成^[36]。

4.2 黄酮及其苷类

黄酮类化合物可以清除氧自由基, 属于强氧化剂, 其具有广泛的药理活性, 如异鼠李素具有心脑血管保护、抗肿瘤、抗炎、抗氧化、预防肥胖等药理作用^[37]。槲皮素可以抗糖尿病、抗肥胖、抗炎、改善循环功能障碍和情绪障碍^[38]。而芦丁具有抗氧化、抑菌、抗炎、抗肿瘤、降血糖、抗过敏等功效^[39]。金线兰中黄酮、黄酮苷类成分众多, 分离得到的黄酮及其苷类化合物包括8-对羟基苄基槲皮素、槲皮素-3-O-葡萄糖苷、槲皮素-3′-O-葡萄糖苷、异鼠李素^[40]、槲皮素、5, 4′-二羟基-6, 7, 3′-三甲氧基黄酮^[29]、芦丁、水仙苷、山柰酚-7-O-葡萄糖苷、异鼠李素-3-O-葡萄糖苷、山柰酚-3-O-葡萄糖苷、山柰酚-7-O-葡萄糖苷、5-hydroxy-3′, 4′, 7-trimethoxyflavonol-3-O-β-D-rutinosid、quercetin-7-O-β-D-[6′′-O-(trans-feruloyl)]-glucopyranoside^[41]、5, 6, 3′, 4′-tetrahydroxy-7, 5′-trimethoflavonol-3′-O-glucoside、kaempferol-3-O-(6′′-p-coumaroyl)-glucoside、山柰酚、quercetin-3, 4′-dimethylether^[42]、槲皮素-7-O-D-葡萄糖苷、槲皮素3-O-β-D-芸香糖苷、异鼠李素-3, 4′-O-β-D-二葡萄糖苷、异鼠李素-3, 7-O-β-D-二葡萄糖苷、异鼠李素-7-O-β-D-二葡萄糖苷^[43]、roxburoside^[30]、鼠李秦素、鼠李秦素-3-O-葡萄糖苷^[31]、山柰酚-7-O-吡喃葡萄糖-(1-3)-吡喃葡萄糖苷、异鼠李素-3-O-α-L-鼠李糖基-(1-6)-β-D-吡喃葡萄糖-(1-3)-吡喃葡萄糖苷^[32], 部分特征性化学成分结构见图 1E。

4.3 有机酸及其糖苷

从金线兰中分离出有机酸及挥发性物质(部分结构见图 1C) 包括软脂酸、硬脂酸、琥珀酸^[28]、香豆酸^[40]、阿魏酸^[29]、棕榈酸^[44]、4-羟基肉桂酸、3-羟基肉桂酸、2-羟基肉桂酸、5-hydroxyferulic acid^[42]、十六烷酸-2, 3-二羟基丙酯^[31]、丁香酸-4-O-葡萄糖苷、Z-香豆酸-4-O-β-D-吡喃葡萄糖苷、Z-阿魏酸-4-O-β-D-吡喃葡萄糖苷。其中代表性成分, 如琥珀酸在抑菌、抗惊厥、镇痛等方面具有显著疗效; 香豆酸在抗血小板活化、抗肿瘤、免疫抑制等方面显示出明显的药理作用; 阿魏酸同样具有抗氧化、抗病毒、抗菌活性, 还可以抗辐射^[44]。

4.4 三萜及甾体类

三萜是一类基本母核由30个碳原子所组成的萜类化合物, 以游离形式或以与糖结合成苷或酯的形式存在于植物体内。三萜类化合物具有溶血、抗癌、抗炎、抗菌、抗病毒、降低胆固醇等活性^[45]。从金线兰中共分离出三萜类化合物7种(部分结构见图 1B), 包括高粱醇、木栓酮^[46]、齐墩果酸^[47]、α-香树脂醇、β-香树脂醇^[31]、sorghumol-3-O-Z-p-coumarate、sorghumol-3-O-E-p-coumarate^[48]。甾体类化合物以环戊烷多氢菲为母核, 具有免疫调节、止血、抗肿瘤、改善心血管疾病等功效^[49]。金线兰中分离出甾体类化合物7种(部分结构见图 1B), 包括胡萝卜苷^[29]、菜油甾醇、β-谷甾醇、豆甾醇、24-异丙烯基胆甾醇、开唇兰甾醇^[46]、羊毛甾醇^[48]。

4.5 多糖类

多糖是由超过10个的单糖通过糖苷键连接形成, 根据在生物体内功能和单糖组成可对多糖种类进行划分。研究发现, 金线兰整株总多糖18%, 根多糖最高可达19%。金线兰与台湾银线兰根多糖与叶中多糖均可以降低链脲佐菌素诱导的糖尿病小鼠的体重、血糖浓度、总胆固醇、低密度脂蛋白胆固醇等一系列糖尿病参数, 且根多糖的活性显著高于叶多糖, 金线兰多糖活性好于台湾银线兰, 提示金线兰多糖具有开发成降血糖药物或功能性食品的潜质^[50]。还有研究表明, 金线兰总多糖对CCl₄诱导的小鼠肝毒性具有一定的保护作用, 可以显著降低血清谷丙转氨酶和谷草转氨酶活性, 降低肝组织丙二醛水平, 提高肝组织的谷胱甘肽水平并增加抗氧化酶的活性, 该作用可能与降低氧化应激、炎症和细胞凋亡有关^[51]。此外, 还可能是通过抑制p38-MAP激酶级联反应, 从而减轻炎症反应和高糖引起的肾损害^[52]。金线兰总多糖还具有抗关节炎作用, 具体表现在可以降低关节炎大鼠的炎症细胞浸润程度及滑膜组织损伤程度, 通过抑制IκB和p65的磷酸化水平来抑制NF-κB通路中相关基因表达, 从而抑制RAW264.7细胞一氧化氮的产生, 最终发挥抗氧化功能^[53]。

4.6 其他化合物

此外, 还从金线兰中检测出十多种核苷类物质, 其中腺苷、尿苷、鸟苷、胞苷含量最高, 几乎不含肌苷。核苷作为核酸及核苷酸的组成成分, 由嘌呤或嘧啶碱与核糖或脱氧核糖缩合而成, 除了具有维持生命活动等基本生理功能外, 部分还可用于治疗恶性肿瘤、风湿性心脏病、肝炎等。含氮化合物主要是指氨基酸及生物碱类化合物。从金线兰中分离出含氮类化合物包括anoectochine^[51]、烟酸、葫芦巴碱(部分结构见图 1D)^[32]。含氮类化合物主要作为药物中间体, 如烟酸, 可合成尼可刹米和烟酸肌醇酯, 用于防治皮肤病和类似的维生素缺乏症, 具有扩张血管的作用, 用于医治末梢神经痉挛、动脉硬化等病症。

5 繁育技术

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金线兰在降血糖、抗肝癌、抗炎等方面具有良好功效, 近年来, 随着金线兰产业规模不断扩大, 市场价格逐年升高, 作为保健品及中药风靡福建省、台湾省, 成为我国发展较快的中药材^[54]。过去的过度开采与不重视, 加之自然竞争力差, 而且金线兰种子必须依靠与真菌共生才可萌发, 且萌发率低, 造成野生金线兰资源稀缺, 供不应求。因此, 目前市场上流通的金线兰主要为人工培养无性繁殖苗。过去由于技术受限, 金线兰始终无法实现人工繁育, 随着无菌栽培与组织培养等技术的发展, 成功解决了金线兰人工繁育的问题。金线兰人工繁育技术主要包括人工培养基培养、设施栽培、仿野生栽培以及菌根栽培。

5.1 人工培养基培养

人工培养基培养是指利用无菌技术将植物在人工配制的富含营养物质的培养基中繁殖的技术。金线兰的人工培养基培养根据是否存在两性生殖细胞结合为受精卵, 可分为无性繁殖培养和有性繁殖培养。无性繁殖通常选择灭菌的幼嫩茎段、茎尖或原球茎作为无性繁殖的外植体, 此方法将外植体诱导成健壮的成苗往往只需要4~5个月时间, 是金线兰产业化最主要的手段^{[55, 56]}。虽然金线兰的无性繁殖技术已相当成熟, 但大规模无性繁殖后, 出现种植退化严重、抗逆性差、化学成分减少等问题。相较于无性繁殖技术, 采用种子繁殖的有性繁殖技术具有繁殖数量大、实生苗抗逆性强、存活率高等特点, 因此, 采用有性繁殖技术对其资源进行优化显得尤为重要。金线兰有性繁殖目前仅在实验室中进行, 并未实现大规模生产, 主要原因在于有性繁殖存在种子萌发率低、萌发时间过长等问题^[57]。

人工培养基培养的成功与否主要与两个因素有关: ①外植体或种子的消毒方式。不同的外植体或种子的消毒条件不同, 一般以75%乙醇浸泡30 s + 0.1%升汞浸泡8~16 min^{[58, 59]}。也有研究发现, 在升汞中加入Tween可以明显降低污染率^[60]。②适宜的培养基。通过筛选不同外植体及发育阶段的培养基, 得到最佳培养基配方为: MS + 1.0 mg·L^-1 6-BA + 1.5 mg·L^-1 NAA + 150 g·L^-1土豆+ 3%蔗糖+ 0.65%琼脂+ 0.15%活性炭^[61]。对不同种子萌发培养基比较后发现, 金线兰种子在1/2 MS培养基上萌发最快。0.3 mg·L^-1生长素NAA对种子萌发有明显的促进作用, 但培养基中细胞分裂素6-BA、ZT和KT对种子萌发的作用不明显, 添加香蕉汁或马铃薯对壮苗具有显著促进效果^[62]。

5.2 设施栽培及仿野生栽培

设施栽培是指利用人工技术手段, 搭建可调控光照、温度等条件的设施, 从而优化植物生长环境, 使植物高效生长的一种栽培技术, 常见的设施如日光薄膜温室、塑料大棚等。金线兰的设施栽培包括单筐套袋式栽培、简易大棚栽培、玻璃温室大棚栽培和连栋钢管大棚栽培。其中, 单筐套袋式栽培成本较高, 工作量较大; 简易大棚栽培成本最低, 但设施功能简陋, 环境调控功能差; 玻璃温室大棚栽培和连栋钢管大棚栽培均可较好地实现种植环境可控, 但成本较高^[54]。总体而言, 设施栽培可初步满足金线兰严格的温、湿度要求, 但温室在温度、光照、湿度可控调节上仍具较大局限, 导致生长期长、存活率低等问题仍待解决。

林下仿野生栽培是指以野生金线兰生长于常绿阔叶林、竹林下的特征为指导, 利用林下特殊的自然环境, 模拟其野生生长环境, 从而促进人工培养金线兰生长的栽培方式。根据栽培模式的不同分为林下小拱棚栽培、林下床式栽培、林下育苗盘栽培和林下梯田栽培。林下仿野生栽培一般选择在具有野生金线兰生长的地方, 通常满足以下条件: 1月份平均气温 < 10 ℃, 7月份平均气温 < 27 ℃, 空气相对湿度 > 70%, 风小但又通风透光, 透光度为30%, 土壤中性或微酸性, pH值4.5~6.5^{[63, 64]}。由于林下仿野生栽培利用自然条件的优势, 可减少生产投入与种植成本, 并且可在理论程度上提高种植金线兰与野生金线兰的一致性, 但是非无菌环境也增加了病虫害的风险。

5.3 菌根栽培

金线兰的种子细小如尘, 每粒蒴果中包含数以万计种子, 但种子发育不完全, 仅具尚未分化的原胚, 自然条件下必须依靠与适宜真菌共生而萌发; 同时, 自然条件下野生金线兰也必须依靠真菌才能完成其完整生长史^[65]。菌根栽培即是利用从植物中分离得到的共生真菌, 建立真菌与金线兰的共生体系, 从而促进其种子萌发、幼苗生长的栽培技术。研究表明, 野生金线兰菌根共生真菌的分离率可达34.38%, 且均形成了典型的菌根结构^[66]。此外, 从兰科其他植物如铁皮石斛中也分离出可与金线兰共生的真菌。共生真菌根据其功能分为促萌发菌和促生长菌, 促萌发菌主要包括亡革菌属Thanatephorus、角担菌Ceratobasidium、小菇属Mycena的开唇兰小菇M. anoectochila、丝核菌属Rhizoctonia、粘帚霉属Gliocladium和瘤菌根菌属Epulorhiza等^[67]。促生长菌株主要来自子囊菌门Ascomycota和担子菌门Basidiomycota, 如毛壳菌属Chaetomium的球毛壳菌C. globosum、炭疽菌属Colletotrichum的炭疽菌C. gloeosporioides, 以及小菇属的兰小菇M. orchidicola和石斛小菇M. dendrobii等^{[68, 69]}。

菌根栽培技术的应用不仅可以显著提高金线兰无性繁殖与有性繁殖的成功率与产量, 研究发现, 该技术还同时促进金线兰中多种活性物质的积累, 提高其药理活性。Yu^[70]通过将分离出的AR-15和AR-18菌根真菌分别与金线兰试管苗共生, 发现除可显著提高鲜重及苗高, 对果糖、蔗糖、多糖及可溶性总糖含量均有不同程度的提高, 且增加了对铬和铁元素的吸收。王涵等^[71]从南平金线兰根中分离出炭垫属菌株cw-10, 发现可促进株高及黄酮、多糖在植物中的积累。Ye等^[68]从金线兰及血叶兰中分离出的球毛壳菌属菌株J162和胶孢炭疽菌属菌株J221不仅可以显著提高金线兰试管苗的生物量, 还促进黄酮、金线莲苷及多糖的积累。研究发现, 共生真菌促生、促萌发机制可能与促进相关基因表达、提高酶活相关。张盈等^{[72, 73]}通过转录组与代谢组联合分析发现, 从金线兰中和分离出的AR-2菌与金线兰共生后可通过提高黄酮相关基因的表达量, 进而显著增加不同时期金线兰中黄酮的积累。刘思思等^[74]通过比较转录组学分析, 发现与金线兰共生后真菌可通过调控GA-GID1-DELLA通路相关基因的表达水平, 进而发挥对金线兰种子的促萌发作用, 为共生关系及其机制提供了新的研究思路。

6 金线兰保护策略及发展建议

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6.1 明确野生资源现状与动态变化

金线兰分布于我国长江以南十余个省份, 其中仅有半数省份对其进行了资源调查, 其余省份野生资源情况仍未知。因此, 首先需要明确各省金线兰的野生资源现状, 包括其分布范围、种群数量、地理位置等, 然后通过定期对代表性地点居群进行样方调查, 了解当地金线兰资源动态变化, 最终建立金线兰全国种质库, 对金线兰标本进行保藏与繁育。

6.2 保护金线兰野生资源

随着人类活动范围的增大, 导致部分低海拔地区的野生金线兰居群环境发生改变, 造成物种消失。同时, 近些年随着金线兰价格提升迅速, 当地居民逐渐意识到金线兰的市场价值, 因此金线兰作为连年被采挖的植物, 野生资源陷入匮乏阶段。建立金线兰自然生态保护区, 保存金线兰及其周边生态系统, 为开展生物科学研究提供良好的基地。同时提高当地居民对金线兰生态保护意识, 加强自然保护区监管力度, 遏制破坏野生金线兰资源的行为。

6.3 完善金线兰人工繁育体系

通过无性、有性等多途径对金线兰进行人工繁育研究, 完善金线兰人工繁育体系。金线兰无性繁殖技术已较成熟, 但活性成分是否改变、种质是否退化等问题亟待明确与验证。因此, 为获得优质金线兰, 对现有繁育技术的完善迫在眉睫。另一方面, 有性繁殖困难急需突破, 主要围绕种子萌发率低, 萌发时间过长等问题。同时, 通过筛选优良种质及促生菌株, 力求尽快将有性繁殖技术融入现有人工繁育体系内, 实现产业技术升级, 达到优质高产。

6.4 开展高质量基础研究

基础研究的突破可以对实际应用提供重要参考与启发。因此, 金线兰基础研究应予以重视。主要包括四方面内容: ①我国开唇兰属植物资源考察; ②金线兰有性繁殖技术及野生抚育; ③金线兰生长发育的分子机制; ④金线兰药效物质基础。

6.5 建立金线兰国家标准与地方标准

基于基础研究与人工繁育技术的发展, 实现金线兰优质高产, 并建立金线兰栽培技术标准。从源头控制与提高金线兰药材质量, 提高金线兰标准化栽培水平。建立金线兰质量评价标准, 保证金线兰药材品质均一稳定与优质高产规模化生产, 促进金线兰产业可持续发展。

7 小结

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本文从我国金线兰的本草考证、资源分布、资源鉴定、化学成分特征与药理活性、繁育技术五方面进行综述, 旨在为金线兰种质资源保护提供参考, 促进金线兰质量标准的建立与统一, 进而推动金线兰产业的可持续发展。作为名贵的兰科药用植物, 金线兰降血糖、抗炎等药效显著, 但其野生资源不足, 人工培养基组培苗是目前市场主要流通品种。而组培苗由于栽培环境及培养条件差异, 导致种质退化问题严重, 并且与野生金线兰存在化学成分差异, 且药效物质不明, 因此人工品种能否完全替代野生金线兰, 仍待进一步研究。同时, 不同地区金线兰种质资源差异较大, 导致药效物质含量及药效参差不齐。自然条件下, 金线兰与真菌共生, 但其分子机制尚未明确, 且不同促生菌的作用方式及作用时间不同, 甚至存在多种促生菌协同作用, 给该问题的研究带来更大挑战。

因此, 明确金线兰生长发育特点及营养需求, 进而筛选出优良种质及促生菌株, 以便快速开展有性繁殖技术研究, 建立菌根栽培体系。并强化种植产区的环境保护, 从而更好将有性繁殖技术及菌根栽培技术应用到实际生产中。在此基础上, 实现金线兰资源可持续利用以及优质高产规模化生产, 为国家大健康产业发展提供物质基础。

作者贡献: 吕欣锴负责文献检索及文章撰写; 周丽思负责文章选题、文献检索及修改文章; 郭顺星负责文章选题、审阅文章。

利益冲突: 本文无任何利益冲突问题。

基金

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国家自然科学基金资助项目(81973426)
国家自然科学基金资助项目(81903749)

参考文献

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2022年第57卷第7期

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doi: 10.16438/j.0513-4870.2022-0019

接收时间：2022-01-05
首发时间：2025-12-23
出版时间：2022-07-12

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收稿日期：2022-01-05
修回日期：2022-02-11

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国家自然科学基金资助项目(81973426)

国家自然科学基金资助项目(81903749)

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中国医学科学院、北京协和医学院药用植物研究所, 北京 100193

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*周丽思, E-mail: zls921203@sina.com;

郭顺星, E-mail: sxguo2015@163.com

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2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

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Latin name	Height /cm	Adaxial side	Abaxial side
Anoectochilus roxburghii	8-18	Dark green to blackish purple	Ovate to ovate-orbicular, 1.3-4.5 × 0.8-3.6 cm, apex acute	Pinkish golden reticulate venation	Pale purplish red
Anoectochilus formosanus	11-20	Dark green	Ovate to ovate-orbicular, 2-5 × 1-3.5 cm, apex acute	White venation	Reddish purple
Anoectochilus chapaensis	12-18	Blackish green	Ovate, oblique, 2-5 × 1.5-3.5 cm, apex acute	Golden-red venation	Pale green
Anoectochilus koshunensis	about 20	Dark green	Ovate to ovate-orbicular, 3-4 × 2-3 cm	White venation	Reddish purple
Anoectochilus lanceolatus	15-30	Dark green	Ovate, ovate-lanceolate, or elliptic, 1.5-8 × 1-5 cm, apex acute	Whitish stripes along midvein and 2 lateral veins	Pale green
Anoectochilus elwesii	15-25	Dark purple or blackish green	Ovate to ovate-orbicular, 1.5-5 × 1-3 cm	3 red veins sometimes	Pale red or pale green
Anoectochilus moulmeinensis	20-30	Green	Oblong to narrowly elliptic, 5-8 × 2-3 cm	Whitish and broad stripes along midvein	Greyish green
Anoectochilus zhejiangensis	8-16	Dark greenish purple	Broadly ovate to ovate-orbicular, 0.7-2.6 × 0.6-2.1 cm, margin slightly undulate, apex acute	Golden-red venation	Pale purplish red
Anoectochilus pingbianensis	15-18	Dark green	Ovate, 3-4.5 × 2-3 cm	Pinkish golden reticulate venation	Pale red or pale green
Anoectochilus burmannicus	16-30	Blackish green	Broadly ovate to ovate-elliptic, slightly oblique, 3-8 × 1.8-5 cm, apex acute	Limited bronze venation	Pinkish red
Anoectochilus emeiensis	19-21	Blackish green	Ovate, 3.5-4 × 3-3.2 cm	Pinkish golden reticulate venation	Reddish purple
Anoectochilus yungianus	10-20	Dark green	Ovate, 1.5-5 × 1-3 cm	Pinkish golden reticulate venation	Dark purple

Latin name

Height /cm

Adaxial side

Abaxial side

Color

Structure

Pattern

Color

Anoectochilus roxburghii

8-18

Dark green to blackish purple

Ovate to ovate-orbicular, 1.3-4.5 × 0.8-3.6 cm, apex acute

Pinkish golden reticulate venation

Pale purplish red

Anoectochilus formosanus

11-20

Dark green

Ovate to ovate-orbicular, 2-5 × 1-3.5 cm, apex acute

White venation

Reddish purple

Anoectochilus chapaensis

12-18

Blackish green

Ovate, oblique, 2-5 × 1.5-3.5 cm, apex acute

Golden-red venation

Pale green

Anoectochilus koshunensis

about 20

Dark green

Ovate to ovate-orbicular, 3-4 × 2-3 cm

White venation

Reddish purple

Anoectochilus lanceolatus

15-30

Dark green

Ovate, ovate-lanceolate, or elliptic, 1.5-8 × 1-5 cm, apex acute

Whitish stripes along midvein and 2 lateral veins

Pale green

Anoectochilus elwesii

15-25

Dark purple or blackish green

Ovate to ovate-orbicular, 1.5-5 × 1-3 cm

3 red veins sometimes

Pale red or pale green

Anoectochilus moulmeinensis

20-30

Green

Oblong to narrowly elliptic, 5-8 × 2-3 cm

Whitish and broad stripes along midvein

Greyish green

Anoectochilus zhejiangensis

8-16

Dark greenish purple

Broadly ovate to ovate-orbicular, 0.7-2.6 × 0.6-2.1 cm, margin slightly undulate, apex acute

Golden-red venation

Pale purplish red

Anoectochilus pingbianensis

15-18

Dark green

Ovate, 3-4.5 × 2-3 cm

Pinkish golden reticulate venation

Pale red or pale green

Anoectochilus burmannicus

16-30

Blackish green

Broadly ovate to ovate-elliptic, slightly oblique, 3-8 × 1.8-5 cm, apex acute

Limited bronze venation

Pinkish red

Anoectochilus emeiensis

19-21

Blackish green

Ovate, 3.5-4 × 3-3.2 cm

Pinkish golden reticulate venation

Reddish purple

Anoectochilus yungianus

10-20

Dark green

Ovate, 1.5-5 × 1-3 cm

Pinkish golden reticulate venation

Dark purple

Latin name	Color	Sepal	Lip resupinate	Lip	Calli	Florescence
A. roxburghii	Pinkish white	Pinkish, outer surface pubescent, 1-veined	No	Pinkish white, Y-shaped; epichile 2-lobed, lobes diverging at an acute angle to one another, obovate-oblong to spatulate; spur conic, apex shallowly bilobed	2 irregular calli, fleshy near entrance of spur	August-December
A. formosanus	White	Reddish brown, pubescent on outer surface	Yes	Pale yellowish white, Y-shaped; epichile longitudinally dilated; spur conic, apex shallowly bilobed	2 calli, near the end of spur, plate	October-November
A. chapaensis	White	Pubescent on outer surface, 1-veined	No	White, Y-shaped; epichile 2-lobed, lobes diverging widely, obovate, margin crenulate, apex truncate; spur conic, apex obtuse	2 calli located near the middle of spur	July-August
A. koshunensis	White	Reddish brown, pubescent on outer surface	No	White, Y-shaped; epichile 2-lobed, slightly brown, margin entire or slightly crenulate; spur conic, extending upward beyond the lobes	2 calli, fleshy near entrance of spur	July-October
A. lanceolatus	Yellow	Yellowish green, glabrous, 1-veined	Yes	Golden yellow, Y-shaped; epichile 2-lobed, cuneate-oblong to obovate, margin entire or slightly crenulate, apex truncate	Slender, needle-like, linear, lanceolate, often hooked upward, sessile	June-September
A. elwesii	White	Green or white	Yes	White, Y-shaped; epichile 2-lobed; nearly obovate; margin entire or slightly crenulate, glabrous	A seperate longitudinal fold, concave near square at top	July-August
A. moulmeinensis	Pinkish red	Pinkish red, outer surface pubescent, 1-veined	Yes	T-shaped, margin slightly crenulate; epichile 2-lobed, nearly obovate; short, margin entire	A longitudinal fold; 2 ovoid, short stipe, apically serrated	August-October
A. zhejiangensis	White	Pinkish red, outer surface pubescent	No	White, Y-shaped; epichile 2-lobed, lobes diverging at an acute angle to one another, flabellate, slightly obliquespur conic, slightly curved, apex shallowly bilobed	2 irregular, stalked calli near entrance	July-September
A. pingbianensis	White	Pinkish	Yes	White, T-shaped; extending forward; epichile 2-lobed, inverted triangle; spur conic	2 calli, fleshly, narrowly elliptic, with short stalk	October
A. burmannicus	Yellowish white	Pink, outer surface pubescent, 1-veined	No	Yellow, Y-shaped; epichile 2-lobed, lobes diverging at an acute angle to one another, oblong, oblanceolate, or spatulatespur conic, apex obtuse	2 subelliptic, sessile calli near entrance	September-December
A. emeiensis	White with purplish red	Fuchsia	No	White, Y-shaped, apex shallowly bilobe; epichile 2-lobed, oblong, oblanceolate; spur straight	2 calli, nearly cuneate	August-September
A. yungianus	White	Greyish green with pinkish, outer surface pubescent	No	White, Y-shaped; epichile 2-lobed, inverted triangle; spur conic	2 calli, near the middle of spur	February

Latin name

Color

Sepal

Lip resupinate

Lip

Calli

Florescence

A. roxburghii

Pinkish white

Pinkish, outer surface pubescent, 1-veined

Pinkish white, Y-shaped; epichile 2-lobed, lobes diverging at an acute angle to one another, obovate-oblong to spatulate; spur conic, apex shallowly bilobed

2 irregular calli, fleshy near entrance of spur

August-December

A. formosanus

White

Reddish brown, pubescent on outer surface

Yes

Pale yellowish white, Y-shaped; epichile longitudinally dilated; spur conic, apex shallowly bilobed

2 calli, near the end of spur, plate

October-November

A. chapaensis

White

Pubescent on outer surface, 1-veined

White, Y-shaped; epichile 2-lobed, lobes diverging widely, obovate, margin crenulate, apex truncate; spur conic, apex obtuse

2 calli located near the middle of spur

July-August

A. koshunensis

White

Reddish brown, pubescent on outer surface

White, Y-shaped; epichile 2-lobed, slightly brown, margin entire or slightly crenulate; spur conic, extending upward beyond the lobes

2 calli, fleshy near entrance of spur

July-October

A. lanceolatus

Yellow

Yellowish green, glabrous, 1-veined

Yes

Golden yellow, Y-shaped; epichile 2-lobed, cuneate-oblong to obovate, margin entire or slightly crenulate, apex truncate

Slender, needle-like, linear, lanceolate, often hooked upward, sessile

June-September

A. elwesii

White

Green or white

Yes

White, Y-shaped; epichile 2-lobed; nearly obovate; margin entire or slightly crenulate, glabrous

A seperate longitudinal fold, concave near square at top

July-August

A. moulmeinensis

Pinkish red

Pinkish red, outer surface pubescent, 1-veined

Yes

T-shaped, margin slightly crenulate; epichile 2-lobed, nearly obovate; short, margin entire

A longitudinal fold; 2 ovoid, short stipe, apically serrated

August-October

A. zhejiangensis

White

Pinkish red, outer surface pubescent

White, Y-shaped; epichile 2-lobed, lobes diverging at an acute angle to one another, flabellate, slightly obliquespur conic, slightly curved, apex shallowly bilobed

2 irregular, stalked calli near entrance

July-September

A. pingbianensis

White

Pinkish

Yes

White, T-shaped; extending forward; epichile 2-lobed, inverted triangle; spur conic

2 calli, fleshly, narrowly elliptic, with short stalk

October

A. burmannicus

Yellowish white

Pink, outer surface pubescent, 1-veined

Yellow, Y-shaped; epichile 2-lobed, lobes diverging at an acute angle to one another, oblong, oblanceolate, or spatulatespur conic, apex obtuse

2 subelliptic, sessile calli near entrance

September-December

A. emeiensis

White with purplish red

Fuchsia

White, Y-shaped, apex shallowly bilobe; epichile 2-lobed, oblong, oblanceolate; spur straight

2 calli, nearly cuneate

August-September

A. yungianus

White

Greyish green with pinkish, outer surface pubescent

White, Y-shaped; epichile 2-lobed, inverted triangle; spur conic

2 calli, near the middle of spur

February