兰科植物原球茎和类原球茎研究进展

doi:10.3969/j.issn.1004-1524.20240581

浙江农业学报 ›› 2025, Vol. 37 ›› Issue (6): 1372-1389.DOI: 10.3969/j.issn.1004-1524.20240581

兰科植物原球茎和类原球茎研究进展

邹俊燕¹^,²(), 王筠竹², 赵婉秋², 尹志浩², 杜建科³^,^*(), 孙崇波²^,^*()

1.中国计量大学生命科学学院,浙江杭州 310018
2.浙江省农业科学院园艺研究所,中药材创新中心,浙江杭州 310021
3.江苏省中国科学院植物研究所(南京中山植物园) 江苏省植物资源保护与利用重点实验室,江苏南京 210014

收稿日期:2024-07-04 出版日期:2025-06-25 发布日期:2025-07-08
作者简介:邹俊燕(1998—),女,河南南阳人,硕士研究生,研究方向为药用植物生殖生长发育。E-mail:3467724827@qq.com
通讯作者: *杜建科,E-mail:15295595190@163.com;孙崇波,E-mail:chongpo1230@163.com
基金资助:
国家自然科学基金青年科学基金(32401644);浙江省自然科学基金(LQ24C160011);浙江省”领雁”研发攻关计划(2023C02028-1);浙江省农业新品种选育重大科技专项(2021C02071-5)

Research progress of protocorm and protocorm-like body in Orchidaceae plants

ZOU Junyan¹^,²(), WANG Yunzhu², ZHAO Wanqiu², YIN Zhihao², DU Jianke³^,^*(), SUN Chongbo²^,^*()

1. College of Life Sciences, China Jiliang University, Hangzhou 310018, China
2. Innovation Center of Chinese Medicine Crops, Institute of Horticulture, Zhejiang Academy of Agricultural Sciences, Hangzhou 310021, China
3. Jiangsu Key Laboratory for Conservation and Utilization of Plant Resources, Institute of Botany, Jiangsu Province and Chinese Academy of Sciences (Nanjing Botanical Garden Men. Sun Yat-sen), Nanjing 210014, China

Received:2024-07-04 Online:2025-06-25 Published:2025-07-08

摘要/Abstract

摘要：

原球茎和类原球茎作为兰科植物生长发育过程中的重要中间繁殖体,不仅是重要的再生器官,部分种属还含有丰富的药用活性成分,具有重要的科研价值与应用潜力。然而,目前对其发生机制的认识仍不够深入,在产业化开发与资源化利用方面也存在明显局限。本文系统综述了原球茎和类原球茎在兰科植物育种、种苗繁育和资源开发中的重要作用,重点从形态建成与发育过程的细胞学特征、内源与环境影响因素、分子调控机制、开发利用现状等方面进行综述。针对当前研究中存在的发生机理不明、技术开发不足等关键问题,本文进行了深入探讨,并展望了其在种苗规模化生产、新种质创制与高附加值产品开发等领域的应用前景,以期为兰科植物原球茎和类原球茎的深入研究与综合利用提供理论参考和实践指导。

关键词: 兰科, 原球茎, 类原球茎, 生长发育, 开发利用

Abstract:

Protocorm and protocorm-like body (PLB) serve as important intermediate propagules during the growth and development of orchid plants. They are not only important regenerative organs but also rich in bioactive compounds in certain species, demonstrating significant scientific value and application potential. However, current understanding of their formation mechanisms remains limited, and there are notable constraints in industrial development and resource utilization. This paper systematically reviews the vital roles of protocorm and PLB in orchid breeding, seedling propagation, and resource exploitation, with a focus on the following aspects: cytological characteristics during morphogenesis and development, endogenous and environmental influencing factors, molecular regulatory mechanisms, and current exploitation and utilization status. Addressing key challenges such as unclear formation mechanisms and insufficient technological development, this study provides an in-depth discussion and prospects their future applications in large-scale seedling production, novel germplasm creation, and high-value product development. The findings aim to offer theoretical references and practical guidance for further research and comprehensive utilization of orchid protocorm and PLB.

Key words: Orchidaceae, protocorm, protocorm-like body, growth and development, development and utilization

中图分类号:

S682.31

邹俊燕, 王筠竹, 赵婉秋, 尹志浩, 杜建科, 孙崇波. 兰科植物原球茎和类原球茎研究进展[J]. 浙江农业学报, 2025, 37(6): 1372-1389.

ZOU Junyan, WANG Yunzhu, ZHAO Wanqiu, YIN Zhihao, DU Jianke, SUN Chongbo. Research progress of protocorm and protocorm-like body in Orchidaceae plants[J]. Acta Agriculturae Zhejiangensis, 2025, 37(6): 1372-1389.

图/表 2

参考文献 120

[1]	LI Y, ZHANG B, YU H. Molecular genetic insights into orchid reproductive development[J]. Journal of Experimental Botany, 2022, 73(7): 1841-1852.
[2]	ZHANG D Y, ZHAO X W, LI Y Y, et al. Advances and prospects of orchid research and industrialization[J]. Horticulture Research, 2022, 9: uhac220.
[3]	YEUNG E C. A perspective on orchid seed and protocorm development[J]. Botanical Studies, 2017, 58(1): 33.
[4]	CARDOSO J C, ZANELLO C A, CHEN J T. An overview of orchid protocorm-like bodies: mass propagation, biotechnology, molecular aspects, and breeding[J]. International Journal of Molecular Sciences, 2020, 21(3): 985.
[5]	LI C R, DONG N, ZHAO Y M, et al. A review for the breeding of orchids: current achievements and prospects[J]. Horticultural Plant Journal, 2021, 7(5): 380-392.
[6]	孙崇波, 刘玫, 谢鸣, 等. 大花蕙兰类原球茎薄切片高频诱导体系的建立[J]. 浙江农业学报, 2008, 20(5): 313-317.
	SUN C B, LIU M, XIE M, et al. Establishment of highly regeneration system in vitro of protocorn-like body thin cell layers of Cymbidium hybridum[J]. Acta Agriculturae Zhejiangensis, 2008, 20(5): 313-317. (in Chinese with English abstract)
[7]	PAEK K Y, HAHN E J, PARK S Y. Micropropagation of Phalaenopsis orchids via protocorms and protocorm-like bodies[J]. Methods in Molecular Biology, 2011, 710: 293-306.
[8]	TEIXEIRA DA SILVA J A, CARDOSO J C, DOBRÁNSZKI J, et al. Dendrobium micropropagation: a review[J]. Plant Cell Reports, 2015, 34(5): 671-704.
[9]	刘咏. 霍山石斛类原球茎液体培养及其保健功效的研究[D]. 合肥: 合肥工业大学, 2005.
	LIU Y. Liquid culture of protocorm-like bodies from Dendrobium huoshanense and its health care efficacy[D]. Hefei: Hefei University of Technology, 2005. (in Chinese with English abstract)
[10]	徐路. 铁皮石斛原球茎饮料的加工技术研究[D]. 湛江: 广东海洋大学, 2015.
	XU L. Research on processing technology of Dendrobium bulb beverage[D]. Zhanjiang: Guangdong Ocean University, 2015. (in Chinese with English abstract)
[11]	黄晓莉, 韦钰, 张琪, 等. 铁皮石斛类原球茎提取物的护肤功效研究[J]. 广州化工, 2020, 48(4): 68-70.
	HUANG X L, WEI Y, ZHANG Q, et al. Skin protection effect of extracts from Dendrobium officinale protocorm-like bodies[J]. Guangzhou Chemical Industry, 2020, 48(4): 68-70. (in Chinese with English abstract)
[12]	BHATTACHARYYA P, KUMARIA S, TANDON P. High frequency regeneration protocol for Dendrobium nobile: a model tissue culture approach for propagation of medicinally important orchid species[J]. South African Journal of Botany, 2016, 104: 232-243.
[13]	WANG H Z, CHEN X Y, YAN X Y, et al. Induction, proliferation, regeneration and kinsenoside and flavonoid content analysis of the Anoectochilus roxburghii(Wall.) Lindl protocorm-like body[J]. Plants, 2022, 11(19): 2465.
[14]	BATYGINA T B, BRAGINA E A, VASILYEVA V E. The reproductive system and germination in orchids[J]. Acta Biologica Cracoviensia Series Botanica, 2003, 45(2): 21-34.
[15]	孙崇波, 刘玫, 施季森, 等. 蕙兰种子无菌萌发及植株再生[J]. 浙江农业学报, 2008, 20(4): 231-235.
	SUN C B, LIU M, SHI J S, et al. Aseptic germination of Cymbidium faberi seeds and in vitro plant regeneration[J]. Acta Agriculturae Zhejiangensis, 2008, 20(4): 231-235. (in Chinese with English abstract)
[16]	DA SILVA J A T. Orchids: Advances in tissue culture, genetics, phytochemistry and transgenic biotechnology[J]. Floriculture and Ornamental Biotechnology, 2013, 7(1): 1-52.
[17]	ZHAO P, WU F, FENG F S, et al. Protocorm-like body (PLB) formation and plant regeneration from the callus culture of Dendrobium candidum Wall ex Lindl[J]. In Vitro Cellular & Developmental Biology-Plant, 2008, 44(3): 178-185.
[18]	LEE Y I, HSU S T, YEUNG E C. Orchid protocorm-like bodies are somatic embryos[J]. American Journal of Botany, 2013, 100(11): 2121-2131.
[19]	HUANG Y W, TSAI Y J, CHEN F C. Characterization and expression analysis of somatic embryogenesis receptor-like kinase genes from Phalaenopsis[J]. Genetics and Molecular Research, 2014, 13(4): 10690-10703.
[20]	GUO B Y, ZENG S J, YIN Y Y, et al. Characterization of phytohormone and transcriptome profiles during protocorm-like bodies development of Paphiopedilum[J]. BMC Genomics, 2021, 22(1): 806.
[21]	FANG S C, CHEN J C, CHANG P Y, et al. Co-option of the SHOOT MERISTEMLESS network regulates protocorm-like body development in Phalaenopsis aphrodite[J]. Plant Physiology, 2022, 190(1): 127-145.
[22]	张丕方, 董崇楣, 李瑶, 等. 虎头兰组织培养中原球茎的形态发生[J]. 复旦学报(自然科学版), 1989, 28(4): 434-437.
	ZHANG P F, DONG C M, LI Y, et al. Morphogenesis of protocorm in Cymbidium grandiflorum in vitro[J]. Journal of Fudan University, 1989, 28(4): 434-437. (in Chinese with English abstract)
[23]	徐程, 詹忠根, 张铭. 中国兰的组织培养[J]. 植物生理学通讯, 2002, 38(2): 171-174.
	XU C, ZHAN Z G, ZHANG M. Tissue culture of Chinese orchid[J]. Plant Physiology Communications, 2002, 38(2): 171-174. (in Chinese)
[24]	苏莹, 刘珍珍, 李会宁, 等. 白及种子萌发及幼苗的微形态学观察[J]. 陕西理工大学学报(自然科学版), 2018, 34(6): 59-64.
	SU Y, LIU Z Z, LI H N, et al. Micromorphological observation of seed germination and seedling of Bletilla striata[J]. Journal of Shaanxi University of Technology(Natural Science Edition), 2018, 34(6): 59-64. (in Chinese with English abstract)
[25]	张凤, 朱艳, 许娜, 等. 霍山石斛种子萌发及原球茎发育的解剖结构观察[J]. 植物资源与环境学报, 2018, 27(3): 115-117.
	ZHANG F, ZHU Y, XU N, et al. Observation on anatomical structures of seed germination and protocorm development of Dendrobium huoshanense[J]. Journal of Plant Resources and Environment, 2018, 27(3): 115-117. (in Chinese with English abstract)
[26]	PARK S Y, MURTHY H N, PAEK K Y. Protocorm-like body induction and subsequent plant regeneration from root tip cultures of Doritaenopsis[J]. Plant Science, 2003, 164(6): 919-923.
[27]	詹忠根, 徐程, 张铭, 等. 铁皮石斛离体根尖经体细胞胚再生植株研究[J]. 浙江大学学报(农业与生命科学版), 2005, 31(5): 579-580.
	ZHAN Z G, XU C, ZHANG M, et al. Plant regeneration throught somatic embryogenesis on root-tip explants of Dendrobium officinale Kimura et Miga[J]. Journal of Zhejiang University(Agric & Life Sci), 2005, 31(5): 579-580. (in Chinese with English abstract)
[28]	詹忠根. 铁皮石斛根尖诱导丛生芽研究[J]. 中草药, 2006, 37(6): 928-931.
	ZHAN Z G. Cluster shoots induction from root-tip segments of Dendrobium officinale[J]. Chinese Traditional and Herbal Drugs, 2006, 37(6): 928-931. (in Chinese with English abstract)
[29]	金青, 马绍鋆, 蔡永萍, 等. 霍山石斛类原球茎诱导及其发育过程研究[J]. 园艺学报, 2009, 36(10): 1525-1530.
	JIN Q, MA S J, CAI Y P, et al. Induction of protocorm-like bodies and its development from stem segments of Dendrobium huoshanense[J]. Acta Horticulturae Sinica, 2009, 36(10): 1525-1530. (in Chinese with English abstract)
[30]	王燕萍, 王广东, 吴震. 大花蕙兰类原球茎形态发生的组织学研究[J]. 亚热带植物科学, 2008, 37(1): 25-28.
	WANG Y P, WANG G D, WU Z. Anatomy study on protocorm-like bodies formation and development of Cymbidium hybridium[J]. Subtropical Plant Science, 2008, 37(1): 25-28. (in Chinese with English abstract)
[31]	张启香, 付素静, 方炎明, 等. 铁皮石斛拟原球茎的发生过程[J]. 浙江林学院学报, 2009, 26(3): 444-448.
	ZHANG Q X, FU S J, FANG Y M, et al. Protocorm-like body (PLB) development on Dendrobium officinale[J]. Journal of Zhejiang Forestry College, 2009, 26(3): 444-448. (in Chinese with English abstract)
[32]	邹嘉欣. 铁皮石斛原球茎形态发生和NAC家族的生物信息学分析[D]. 成都: 西南交通大学, 2014.
	ZOU J X. Dendrobium candidum Wall ex Lindl protocorm morphogenesis and bioinformatics analysis of NAC family[D]. Chengdu: Southwest Jiaotong University, 2014. (in Chinese with English abstract)
[33]	朱国兵, 杨柏云, 敖爱艳. 寒兰的组织培养与试管开花[J]. 植物生理学通讯, 2008, 44(3): 513-514.
	ZHU G B, YANG B Y, AO A Y. Tissue culture and in vitro flowering of Cymbidium kanran Makino[J]. Plant Physiology Communications, 2008, 44(3): 513-514. (in Chinese with English abstract)
[34]	刘鑫. 春剑种子胚胎发育及非共生萌发研究[D]. 绵阳: 西南科技大学, 2020.
	LIU X. Study on embryo development and non-symbiotic germination of Cymbidium tortisepalum var. longibracteatum[D]. Mianyang: Southwest University of Science and Technology, 2020. (in Chinese with English abstract)
[35]	聂宁, 朱艳, 田梅, 等. 白及种子萌发及原球茎发育过程的细胞组织学观察[J]. 中国中药杂志, 2016, 41(8): 1446-1449.
	NIE N, ZHU Y, TIAN M, et al. Morphological and cytohistological observations of seed germination and protocorm development of Bletilla striata[J]. China Journal of Chinese Materia Medica, 2016, 41(8): 1446-1449. (in Chinese with English abstract)
[36]	CHUGH S, GUHA S, RAO I U. Micropropagation of orchids: a review on the potential of different explants[J]. Scientia Horticulturae, 2009, 122(4): 507-520.
[37]	魏丽芳, 于桂芬, 冯国宝, 等. 铁皮石斛组织培养与快速繁殖研究进展[J]. 安徽农业科学, 2013, 41(35): 13561-13563.
	WEI L F, YU G F, FENG G B, et al. Advances in tissue culture and rapid propagation of Dendrobium candidum Wall. ex Lindl[J]. Journal of Anhui Agricultural Sciences, 2013, 41(35): 13561-13563. (in Chinese with English abstract)
[38]	刘扬, 李宏杨, 任杰, 等. 兜唇石斛种子无菌播种与快速繁殖[J]. 热带农业科学, 2020, 40(5): 34-41.
	LIU Y, LI H Y, REN J, et al. Aseptic seeding and rapid propagation of Dendrobium aphyllum[J]. Chinese Journal of Tropical Agriculture, 2020, 40(5): 34-41. (in Chinese with English abstract)
[39]	符红艳, 黄国林, 周宇霞, 等. 白芨种子萌发条件及其显微结构观察[J/OL]. 分子植物育种, 2022: 1-8. (2022-09-17)[2024-06-24]. https://kns.cnki.net/KCMS/detail/detail.aspxfilename=FZZW2022091500N&dbname=CJFD&dbcode=CJFQ.
	FU H Y, HUANG G L, ZHOU Y X, et al. Observation on germination conditions and microstructure of Bletilla striata seeds[J/OL]. Molecular Plant Breeding, 2022: 1-8. (2022-09-17)[2024-06-24]. https://kns.cnki.net/KCMS/detail/detail.aspxfilename=FZZW2022091500N&db-name=CJFD&dbcode=CJFQ.
[40]	张东旭. 大花蕙兰杂交及种胚萌发研究[D]. 泰安: 山东农业大学, 2010.
	ZHANG D X. Study on cross breeding and aseptic germination of embryo of Cymbidium hybridium[D]. Tai’an: Shandong Agricultural University, 2010. (in Chinese with English abstract)
[41]	李风童, 包建忠, 孙叶, 等. 蕙兰胚珠发育及主要发育事件分布[J]. 核农学报, 2015, 29(12): 2300-2306.
	LI F T, BAO J Z, SUN Y, et al. Ovule development and major developmental events distribution in Cymbidium faberi[J]. Journal of Nuclear Agricultural Sciences, 2015, 29(12): 2300-2306. (in Chinese with English abstract)
[42]	林江波, 王伟英, 李海明, 等. 铁皮石斛茎段原球茎的诱导、分化与植株再生[J]. 福建农业学报, 2016, 31(10): 1075-1079.
	LIN J B, WANG W Y, LI H M, et al. Protocorm induction/differentiation and plantlet regeneration of stems from Dendrobium officinale Kimura et Migo[J]. Fujian Journal of Agricultural Sciences, 2016, 31(10): 1075-1079. (in Chinese with English abstract)
[43]	陈芳, 覃顺旺, 王跃华, 等. 杜鹃兰类原球茎快速增殖研究[J]. 种子, 2021, 40(6): 141-145.
	CHEN F, QIN S W, WANG Y H, et al. Study on rapid proliferation of protocorm-like bodies in Cremastra appendiculata[J]. Seed, 2021, 40(6): 141-145. (in Chinese with English abstract)
[44]	夏科, 吴巧芬, 赵志国, 等. 石斛属植物再生及遗传转化研究进展[J]. 亚热带农业研究, 2020, 16(2): 110-117.
	XIA K, WU Q F, ZHAO Z G, et al. Research progress on plant regeneration and genetic transformation of the Dendrobium genus[J]. Subtropical Agriculture Research, 2020, 16(2): 110-117. (in Chinese with English abstract)
[45]	陈甘牛, 苗杰, 陆兆军, 等. ‘章姬’草莓茎尖脱毒及快繁技术研究[J]. 山东林业科技, 2022, 52(1): 31-35.
	CHEN G N, MIAO J, LU Z J, et al. Study on virus-free and rapid propagation of strawberry stem tip of ‘Zhangji’[J]. Journal of Shandong Forestry Science and Technology, 2022, 52(1): 31-35. (in Chinese with English abstract)
[46]	刘晓燕, 向青云, 刘玲玲, 等. 基本培养基及附加物对蝴蝶兰原球茎增殖效果的影响[J]. 种子, 2005, 24(6): 18-20.
	LIU X Y, XIANG Q Y, LIU L L, et al. The effect of basic culture media and additional compounds on the propagation of phalaenosis PLB[J]. Seed, 2005, 24(6): 18-20. (in Chinese with English abstract)
[47]	任建宏, 乔永旭, 李彦辉, 等. 蝴蝶兰叶片PLB诱导因子的优化[J]. 北方园艺, 2010(22): 139-142.
	REN J H, QIAO Y X, LI Y H, et al. Optimization on factor of PLB induced by leaves of Phalaenopsis[J]. Northern Horticulture, 2010(22): 139-142. (in Chinese with English abstract)
[48]	韩思仪, 崔永一. 蝴蝶兰原球茎组织防褐化研究[J]. 江苏农业科学, 2019, 47(13): 67-71.
	HAN S Y, CUI Y Y. Study on anti-browning of protocorm tissue of Phalaenopsis taisuco tissue[J]. Jiangsu Agricultural Sciences, 2019, 47(13): 67-71. (in Chinese)
[49]	DE MELO FERREIRA W, DE OLIVEIRA A M, VIANA J C, et al. Asymbiotic germination, initial development in vitro and acclimatization of Cyrtopodium paludicolum Hoehne, a Brazilian Savanna orchid species[J]. Rodriguésia, 2022, 73: e01272020.
[50]	李玉萍, 王燕青, 武文婷, 等. 春兰与大花蕙兰杂交种原球茎分化和生根研究[J]. 天津农业科学, 2015, 21(8): 127-132.
	LI Y P, WANG Y Q, WU W T, et al. Study on differentiation and rooting of hybrid protocorm-like bodies of crosses between Cymbidium goeringii and C. hybridium[J]. Tianjin Agricultural Sciences, 2015, 21(8): 127-132. (in Chinese with English abstract)
[51]	付双彬, 池梦薇, 杨燕萍, 等. 虎头兰类原球茎诱导、增殖及保存[J]. 北方园艺, 2020(10): 71-77.
	FU S B, CHI M W, YANG Y P, et al. Induction, propagation and conservation of protocorm-like bodys of Cymbidium hookerianum[J]. Northern Horticulture, 2020(10): 71-77. (in Chinese with English abstract)
[52]	XU C Y, CAO H F, ZHANG Q Q, et al. Control of auxin-induced callus formation by bZIP59-LBD complex in Arabidopsis regeneration[J]. Nature Plants, 2018, 4(2): 108-115.
[53]	UTAMI E S W, HARIYANTO S. Organic compounds: contents and their role in improving seed germination and protocorm development in orchids[J]. International Journal of Agronomy, 2020, 2020(1): 2795108.
[54]	蔡宣梅. 福建铁皮石斛的组培快繁体系的研究[J]. 福建热作科技, 2020, 45(4): 1-4.
	CAI X M. Study on tissue culture and rapid propagation system of Fujian’s Dendrobium candidum Wall. ex Lindl[J]. Fujian Science & Technology of Tropical Crops, 2020, 45(4): 1-4. (in Chinese with English abstract)
[55]	叶睿华, 吕享, 李小兰, 等. 五种抗褐化剂对杜鹃兰原球茎增殖培养的作用效果[J]. 植物生理学报, 2018, 54(6): 1103-1110.
	YE R H, LÜ X, LI X L, et al. Effects of five browning inhibitors on protocorms proliferation culture of Cremastra appendiculata[J]. Plant Physiology Journal, 2018, 54(6): 1103-1110. (in Chinese with English abstract)
[56]	吴彦秋, 吕享, 李小兰, 等. 杜鹃兰原球茎增殖培养条件[J]. 北方园艺, 2016(19): 124-128.
	WU Y Q, LYU X, LI X L, et al. Culture conditions of protocorms proliferation of Cremastra appendiculata[J]. Northern Horticulture, 2016(19): 124-128. (in Chinese with English abstract)
[57]	郑晓君, 叶静, 管常东, 等. 兰科植物种子萌发研究进展[J]. 北方园艺, 2010(19): 206-209.
	ZHENG X J, YE J, GUAN C D, et al. Review on recent progress of orchid seeds germination[J]. Northern Horticulture, 2010(19): 206-209. (in Chinese with English abstract)
[58]	杨建文, 凌鸿, 张盈, 等. 内生真菌对兰科药用植物种子萌发作用研究进展[J]. 菌物学报, 2018, 37(1): 22-34.
	YANG J W, LING H, ZHANG Y, et al. Effects of endophytic fungi on seed germination of medicinal plants of Orchidaceae: a review[J]. Mycosystema, 2018, 37(1): 22-34. (in Chinese with English abstract)
[59]	高越. 濒危兰科药用植物手参(Gymnadenia conopsea)种子的真菌共生萌发研究[D]. 北京: 北京协和医学院, 2020.
	GAO Y. Study on Fungal symbiosis germination of endangered orchidaceae medicinal plant (Gymnadenia conopsea)[D]. Beijing: Peking Union Medical College, 2020. (in Chinese with English abstract)
[60]	许凤来, 朱志炎, 何勇, 等. 印度梨形孢对铁皮石斛种子萌发和原球茎生长的影响[J]. 热带亚热带植物学报, 2021, 29(1): 59-66.
	XU F L, ZHU Z Y, HE Y, et al. Effects of Piriformospora indica on seed germination and protocorm growth of Dendrobium officinale[J]. Journal of Tropical and Subtropical Botany, 2021, 29(1): 59-66. (in Chinese with English abstract)
[61]	OLIYA B K, CHAND K, THAKURI L S, et al. Assessment of genetic stability of micropropagated plants of Rhynchostylis retusa(L.) using RAPD markers[J]. Scientia Horticulturae, 2021, 281: 110008.
[62]	徐步青, 崔永一, 郭岑, 等. 不同光照强度和培养时间下铁皮石斛类原球茎生物量、多糖和生物碱量的动态变化[J]. 中草药, 2012, 43(2): 355-359.
	XU B Q, CUI Y Y, GUO C, et al. Dynamic variation of biomass and content of polysaccharide and alkaloid in protocorm like bodies from Dendrobium officinale at different light intensities and incubation time[J]. Chinese Traditional and Herbal Drugs, 2012, 43(2): 355-359. (in Chinese with English abstract)
[63]	AHMADI CHASHMI K, OMRAN V O G, EBRAHIMI R, et al. Light quality affects protocorm-like body (PLB) formation, growth and development of in vitro plantlets of Phalaenopsis pulcherrima[J]. Biology Bulletin, 2022, 49(5): 476-484.
[64]	刘敏, 舒雨婷, 张玉琼, 等. 白芨种子萌发及其原球茎高效增殖体系建立[J]. 南方农业学报, 2017, 48(12): 2223-2227.
	LIU M, SHU Y T, ZHANG Y Q, et al. Seed germination and effective protocorm proliferation system establishment for Bletilla striata(Thumb.) Reichb.f[J]. Journal of Southern Agriculture, 2017, 48(12): 2223-2227. (in Chinese with English abstract)
[65]	陈青青, 王丛巧, 赖钟雄. 温差对铁皮石斛原球茎生长及药用成分的影响[J]. 园艺与种苗, 2022, 42(2): 20-23.
	CHEN Q Q, WANG C Q, LAI Z X. Effects of temperature difference on PLBs growth and medicinal components of Dendrobium officinale[J]. Horticulture & Seed, 2022, 42(2): 20-23. (in Chinese with English abstract)
[66]	朱星扬, 韩静, 朱洁莹, 等. 磷源对铁皮石斛原球茎中多糖积累的影响[J]. 科技通报, 2020, 36(5): 71-76.
	ZHU X Y, HAN J, ZHU J Y, et al. Effects of phosphorus on accumulation of polysaccharides in protocorm of Dendrobium officinale[J]. Bulletin of Science and Technology, 2020, 36(5): 71-76. (in Chinese with English abstract)
[67]	WEI M, YANG C Y, WEI S H. Enhancement of the differentiation of protocorm-like bodies of Dendrobium officinale to shoots by ultrasound treatment[J]. Journal of Plant Physiology, 2012, 169(8): 770-774.
[68]	孙晓慧, 姜成平, 赵艳, 等. 中国兰原球茎诱导及组培快繁技术研究[J]. 农业科技通讯, 2023(4): 120-123.
	SUN X H, JIANG C P, ZHAO Y, et al. Study on protocorm induction and tissue culture and rapid propagation of China orchid[J]. Bulletin of Agricultural Science and Technology, 2023(4): 120-123. (in Chinese)
[69]	范静. 铁皮石斛原球茎转录组测序分析[D]. 成都: 西南交通大学, 2018.
	FAN J. Transcriptomic analysis of the protocorms in Dendrobium officinale Kimura et Migo[D]. Chengdu: Southwest Jiaotong University, 2018. (in Chinese with English abstract)
[70]	胡睿, 郭建秀, 郭小强, 等. 铁皮石斛DoTIFY基因家族全基因组鉴定及在原球茎发育过程中的表达[J]. 生物学杂志, 2021, 38(5): 53-58.
	HU R, GUO J X, GUO X Q, et al. Genome-wide identification and analysis of the TIFY gene family in Dendrobium officinale Kimura et Migo during protocorm development[J]. Journal of Biology, 2021, 38(5): 53-58. (in Chinese with English abstract)
[71]	ZHANG G, LI Y M, HU B X, et al. Isolation and differential expression of a novel MAP kinase gene DoMPK4 in Dendrobium officinale[J]. Acta Pharmaceutica Sinica, 2014, 49(7): 1076-1083. (in Chinese with English abstract)
[72]	安红强, 范静, 梁易, 等. 铁皮石斛泛素结合酶基因DoUBC24的克隆及表达分析[J]. 生物技术通讯, 2016, 27(5): 643-648.
	AN H Q, FAN J, LIANG Y, et al. Cloning and expression analysis of ubiquitin-conjugating enzyme 24 in Dendrobium officinale Kimura et Migo[J]. Letters in Biotechnology, 2016, 27(5): 643-648. (in Chinese with English abstract)
[73]	安红强. 铁皮石斛原球茎内参基因的筛选及LEA基因家族表达分析[D]. 成都: 西南交通大学, 2017.
	AN H Q. Selection of internal reference genes in protocorm and the expression analysis of LEA gene family of Dendrobium officinale Kimura et Migo[D]. Chengdu: Southwest Jiaotong University, 2017. (in Chinese with English abstract)
[74]	刘楚琪, 胡睿, 王万军. 铁皮石斛氨基酸通透酶基因(DoAAP2)的克隆及表达分析[J]. 生物学杂志, 2019, 36(6): 6-11.
	LIU C Q, HU R, WANG W J. Cloning and expression analysis of a amino acid permease gene (DoAAP2) in the Dendrobium officinale Kimura et Migo[J]. Journal of Biology, 2019, 36(6): 6-11. (in Chinese with English abstract)
[75]	马凌晖, 欧景丹, 刘荣荣, 等. 铁皮石斛泛素结合酶DoUBC9的克隆鉴定与表达分析[J]. 现代生物医学进展, 2019, 19(13): 2407-2413.
	MA L H, OU J D, LIU R R, et al. Clonal identification and expression analysis of ubiquitin-conjugating enzyme DoUBC9 in Dendrobium officinale Kimura et Migo[J]. Progress in Modern Biomedicine, 2019, 19(13): 2407-2413. (in Chinese with English abstract)
[76]	FANG L, KONG X P, WEN Y T, et al. Characterization of embryo and protocorm development of Paphiopedilum spicerianum[J]. Plant Physiology and Biochemistry, 2021, 167: 1024-1034.
[77]	阎波, 刘思思, 陈娟, 等. 药用植物铁皮石斛ABC转运蛋白基因的鉴定及其差异表达分析[J]. 药学学报, 2018, 53(7): 1177-1189.
	YAN B, LIU S S, CHEN J, et al. Identification and differential expression analysis of ABC transporter gene from medicinal plant Dendrobium officinale[J]. Acta Pharmaceutica Sinica, 2018, 53(7): 1177-1189. (in Chinese with English abstract)
[78]	JIANG Y T, TIAN M, WANG C X, et al. Transcriptome sequencing and differential gene expression analysis reveal the mechanisms involved in seed germination and protocorm development of Calanthe tsoongiana[J]. Gene, 2021, 772: 145355.
[79]	马庆, 付强, 邹西西, 等. 铁皮石斛DELLA家族基因克隆与生物信息学分析[J]. 贵州中医药大学学报, 2022, 44(6): 13-19.
	MA Q, FU Q, ZOU X X, et al. Cloning and bioinformatics analysis of the DELLA family genes in Dendrobium officinale[J]. Journal of Guizhou University of Traditional Chinese Medicine, 2022, 44(6): 13-19. (in Chinese)
[80]	刘思思, 张岗, 陈晓梅, 等. 铁皮石斛赤霉素3-氧化酶基因的克隆及表达分析[J]. 中草药, 2016, 47(6): 990-996.
	LIU S S, ZHANG G, CHEN X M, et al. Cloning and expression analysis of gibberellin 3-oxidase gene in Dendrobium officinale[J]. Chinese Traditional and Herbal Drugs, 2016, 47(6): 990-996. (in Chinese with English abstract)
[81]	赵明明, 张岗, 张大为, 等. 铁皮石斛DoMAPK5基因的克隆及表达特征分析[J]. 时珍国医国药, 2016, 27(2): 509-512.
	ZHAO M M, ZHANG G, ZHANG D W, et al. Cloning and expression analysis of DoMAPK5 in Dendrobium officinale[J]. Lishizhen Medicine and Materia Medica Research, 2016, 27(2): 509-512. (in Chinese with English abstract)
[82]	LI Y Y, CHEN X M, ZHANG Y, et al. Immunolocalization and changes of hydroxyproline-rich glycoproteins during symbiotic germination of Dendrobium officinale[J]. Frontiers in Plant Science, 2018, 9: 552.
[83]	安红强, 王万军. 铁皮石斛胚胎发生相关基因DoEMB8的克隆及表达分析[J]. 生物学杂志, 2017, 34(4): 1-5.
	AN H Q, WANG W J. Cloning and expression analysis of embryogenesis-associated protein EMB8(DoEMB8) in Dendrobium officinale Kimura et Migo[J]. Journal of Biology, 2017, 34(4): 1-5. (in Chinese with English abstract)
[84]	梁易. 铁皮石斛原球茎发育过程中DoWOX与DoSERK的表达分析[D]. 成都: 西南交通大学, 2018.
	LIANG Y. Expression analysis of DoWOX and DoSERK in Dendrobium officinale during protocorm development[D]. Chengdu: Southwest Jiaotong University, 2018. (in Chinese with English abstract)
[85]	FANG S C, CHEN J C, WEI M J. Protocorms and protocorm-like bodies are molecularly distinct from zygotic embryonic tissues in Phalaenopsis aphrodite[J]. Plant Physiology, 2016, 171(4): 2682-2700.
[86]	ZENG D Q, SI C, ZHANG M Z, et al. ERF5 enhances protocorm-like body regeneration via enhancement of STM expression in Dendrobium orchid[J]. Journal of Integrative Plant Biology, 2023, 65(9): 2071-2085.
[87]	DING L, YAN S S, JIANG L, et al. HANABA TARANU regulates the shoot apical meristem and leaf development in cucumber (Cucumis sativus L.)[J]. Journal of Experimental Botany, 2015, 66(22): 7075-7087.
[88]	IWASE A, HARASHIMA H, IKEUCHI M, et al. WIND1 promotes shoot regeneration through transcriptional activation of ENHANCER OF SHOOT REGENERATION1 in Arabidopsis[J]. The Plant Cell, 2017, 29(1): 54-69.
[89]	XU X M, WANG J, XUAN Z Y, et al. Chaperonins facilitate KNOTTED1 cell-to-cell trafficking and stem cell function[J]. Science, 2011, 333(6046): 1141-1144.
[90]	曾丹琦, 张明泽, 何春梅, 等. 铁皮石斛WOX转录因子的鉴定和分析[J]. 热带亚热带植物学报, 2021, 29(3): 301-310.
	ZENG D Q, ZHANG M Z, HE C M, et al. Identification and analysis of WOX transcription factor in Dendrobium officinale[J]. Journal of Tropical and Subtropical Botany, 2021, 29(3): 301-310. (in Chinese with English abstract)
[91]	ZENG D Q, TEIXEIRA DA SILVA J A, ZHANG M Z, et al. Genome-wide identification and analysis of the APETALA2 (AP2) transcription factor in Dendrobium officinale[J]. International Journal of Molecular Sciences, 2021, 22(10): 5221.
[92]	SUDHERSAN C, ABOEL-NIL M, HUSSAIN J. Tissue culture technology for the conservation and propagation of certain native plants[J]. Journal of Arid Environments, 2003, 54(1): 133-147.
[93]	王小乐, 迟天华, 王海滨, 等. 蔗糖和甘露醇复合处理对菊花低温离体保存的影响[J]. 分子植物育种, 2019, 17(5): 1597-1604.
	WANG X L, CHI T H, WANG H B, et al. Effects of sucrose and mannitol combined treatment on in vitro conservation of Chrysanthemum at low temperature[J]. Molecular Plant Breeding, 2019, 17(5): 1597-1604. (in Chinese with English abstract)
[94]	THIERRY C, FLORIN B, PÉTIARD V. Changes in protein metabolism during the acquisition of tolerance to cryopreservation of carrot somatic embryos[J]. Plant Physiology and Biochemistry, 1999, 37(2): 145-154.
[95]	FOLGADO R, SERGEANT K, RENAUT J, et al. Changes in sugar content and proteome of potato in response to cold and dehydration stress and their implications for cryopreservation[J]. Journal of Proteomics, 2014, 98: 99-111.
[96]	MERRITT D J, HAY F R, SWARTS N D, et al. Ex situ conservation and cryopreservation of orchid germplasm[J]. International Journal of Plant Sciences, 2014, 175(1): 46-58.
[97]	VENDRAME W, DE FARIA R T, SORACE M, et al. Orchid cryopreservation[J]. Ciência e Agrotecnologia, 2014, 38(3): 213-229.
[98]	POPOVA E, KIM H H, SAXENA P K, et al. Frozen beauty: the cryobiotechnology of orchid diversity[J]. Biotechnology Advances, 2016, 34(4): 380-403.
[99]	DUBUS F. Emploi du saccharose pur, en mélange avec du glucose ou du glycérol comme cryoprotecteurs pour la congélation de protocormes de Cymbidium en culture in vitro[J]. Bulletin van de Koninklijke Belgische Botanische Vereniging, 1980, 113: 59-62.
[100]	王君晖, 张毅翔, 刘峰, 等. 铁皮石斛种子、原球茎和类原球茎体的超低温保存研究[J]. 园艺学报, 1999, 26(1): 59-61.
	WANG J H, ZHANG Y X, LIU F, et al. Cryopreservation of seeds, protocorms and protocormlikebodies of Dendrobium candidum[J]. Acta Horticulturae Sinica, 1999, 26(1): 59-61. (in Chinese with English abstract)
[101]	王春浩. 文心兰类原球茎玻璃化超低温保存及相关生理生化与组织学研究[D]. 南京: 南京农业大学, 2015.
	WANG C H. The cryopresevation of Oncidium protocorm like bodies by vitrification and their physiological and histological characteristics analysis[D]. Nanjing: Nanjing Agricultural University, 2015. (in Chinese with English abstract)
[102]	张小明, 鲍根良, 叶胜海, 等. 作物人工种子的研究进展[J]. 种子, 2002, 21(2): 41-43.
	ZHANG X M, BAO G L, YE S H, et al. Advances in study on artificial seeds of crop[J]. Seed, 2002, 21(2): 41-43. (in Chinese)
[103]	PRADHAN S, TIRUWA B L, SUBEDEE B R, et al. Efficient plant regeneration of Cymbidium aloifolium(L.) Sw., a threatened orchid of nepal through artificial seed technology[J]. American Journal of Plant Sciences, 2016, 7(14): 1964-1974.
[104]	吴文杰. 文心兰(Oncidium)原球茎液体增殖及人工种子制作研究[D]. 南京: 南京农业大学, 2006.
	WU W J. Establishment of liquid culture systems for protocorm like bodies and artificial seeds making of Oncidium[D]. Nanjing: Nanjing Agricultural University, 2006. (in Chinese with English abstract)
[105]	曾颖苹, 朱乾坤, 王万军. 铁皮石斛人工种子包衣技术研究[J]. 北方园艺, 2012(17): 155-158.
	ZENG Y P, ZHU Q K, WANG W J. The coating on artificial seeds of Dendrobium candidum Wall ex Lindl[J]. Northern Horticulture, 2012(17): 155-158. (in Chinese with English abstract)
[106]	张明生, 彭斯文, 杨小蕊, 等. 杜鹃兰人工种子技术研究[J]. 中国中药杂志, 2009, 34(15): 1894-1897.
	ZHANG M S, PENG S W, YANG X R, et al. Preparation technique of Cremastra appendiculata synthetic seed[J]. China Journal of Chinese Materia Medica, 2009, 34(15): 1894-1897. (in Chinese with English abstract)
[107]	张建彬, 王璐, 彭宵, 等. 大花蕙兰类原球茎的离体培养及人工种子制作与萌发的研究[J]. 种子, 2014, 33(6): 10-13.
	ZHANG J B, WANG L, PENG X, et al. Studies on vitro culture of protocorm-like bodies of Cymbidium hybridum and its artificial seed production and germination[J]. Seed, 2014, 33(6): 10-13. (in Chinese with English abstract)
[108]	MII M, CHIN D P. Genetic transformation on orchid species: an overview of approaches and methodologies[M]// Orchid propagation: from laboratories to greenhouses:methods and protocols. New York: Springer New York, 2018: 347-365.
[109]	MISHIBA K I, CHIN D P, MII M. Agrobacterium-mediated transformation of Phalaenopsis by targeting protocorms at an early stage after germination[J]. Plant Cell Reports, 2005, 24(5): 297-303.
[110]	UTAMI E S W, HARIYANTO S, MANUHARA Y S W. Agrobacterium tumefaciens-mediated transformation of Dendrobium lasianthera J.J.Sm: an important medicinal orchid[J]. Journal of Genetic Engineering and Biotechnology, 2018, 16(2): 703-709.
[111]	KUEHNLE A R, SUGII N. Transformation of Dendrobium orchid using particle bombardment of protocorms[J]. Plant Cell Reports, 1992, 11(9): 484-488.
[112]	YOU S J, LIAU C H, HUANG H E, et al. Sweet pepper ferredoxin-like protein (PFLP) gene as a novel selection marker for orchid transformation[J]. Planta, 2003, 217(1): 60-65.
[113]	CHEW Y C, WAN ABDULLAH W M A N, ANDREW KOK D X, et al. Development of an efficient particle bombardment transformation system for the endemic orchid, Phalaenopsis bellina[J]. Sains Malaysiana, 2019, 48(9): 1867-1877.
[114]	易双双, 陆顺教, 杨光穗, 等. 兰花遗传转化技术研究进展[J]. 北方园艺, 2016(20): 187-194.
	YI S S, LU S J, YANG G S, et al. Research progress on orchids genetic transformation technology[J]. Northern Horticulture, 2016(20): 187-194. (in Chinese with English abstract)
[115]	陈稷, 张浪, 黄进. 兰科植物转基因技术研究进展[J]. 生物学教学, 2022, 47(4): 2-5.
	CHEN J, ZHANG L, HUANG J. Progress in transgenic technology of Orchidaceae[J]. Biology Teaching, 2022, 47(4): 2-5. (in Chinese)
[116]	张成才, 王升, 王月枫, 等. 药用植物组织培养技术在中药资源可持续发展中的应用研究[J]. 中国中药杂志, 2023, 48(5): 1186-1193.
	ZHANG C C, WANG S, WANG Y F, et al. Application of tissue culture technology of medicinal plants in sustainable development of Chinese medicinal resources[J]. China Journal of Chinese Materia Medica, 2023, 48(5): 1186-1193. (in Chinese with English abstract)
[117]	WANG H Q, JIN M Y, PAEK K Y, et al. An efficient strategy for enhancement of bioactive compounds by protocorm-like body culture of Dendrobium candidum[J]. Industrial Crops and Products, 2016, 84: 121-130.
[118]	LIN W, WANG J J, XU X M, et al. Rapid propagation in vitro and accumulation of active substances of endangered Dendrobium cariniferum Rchb. f[J]. Bioengineered, 2020, 11(1): 386-396.
[119]	韦丽文, 辛宁, 李培泰, 等. 铁皮石斛与铁皮石斛原球茎质量比较的研究进展[J]. 中国医药指南, 2013, 11(25): 59-61.
	WEI L W, XIN N, LI P T, et al. Research progress on the quality comparison between Dendrobium officinale and Dendrobium officinale protocorm[J]. Guide of China Medicine, 2013, 11(25): 59-61. (in Chinese)
[120]	王玲, 唐德强, 王佳佳, 等. 铁皮石斛原球茎与野生铁皮石斛多糖的抗菌及体外抗氧化活性比较[J]. 西北农林科技大学学报(自然科学版), 2016, 44(6): 167-172.
	WANG L, TANG D Q, WANG J J, et al. Comparison of antibacterial and in vitro antioxidant activities of polysaccharides from Dendrobium candidum protocorms and wild Dendrobium[J]. Journal of Northwest A&F University(Natural Science Edition), 2016, 44(6): 167-172. (in Chinese with English abstract)

分类 Classification	基因名称 Gene name	物种 Species	鉴定方法 Verification method	功能 Function	参考文献 Reference
原球茎Protocorm
胚胎发育相关基因 Embryonic development related genes	PsSERKs、PsBBMs	白旗兜兰 Paphiopedilum spicerianum	转录组、实时荧光定量PCR(qRT-PCR) Transcriptome analysis、Real-Time PCR(qRT-PCR)	参与原球茎的发育 Involve in the development of protocorm	[76]
SAM形成相关基因 Shoot apical meristem related genes	PsWOX8、PsCLAVATA2、 PsCUC2、PsSCR	白旗兜兰 P. spicerianum	转录组、qRT-PCR Transcriptome analysis、qRT-PCR	参与原球茎中顶端分生组织SAM形成 Involve in shoot apical meristem formation in protocorm	[76]
ABA生物合成基因 ABA biosynthesis genes	CtNCED1、CtZEP	无距虾脊兰 Calanthe tsoongiana	qRT-PCR、激素含量检测 qRT-PCR, detection of endogenous hormone content	参与原球茎发育 Involve in the development of protocorm	[78]
IAA合成关键酶基因 IAA synthesis key enzymes gene	CtTSA1	无距虾脊兰 C. tsoongiana	qRT-PCR、激素含量检测 qRT-PCR, detection of endogenous hormone content	在原球茎发育和分化中具有潜在作用 An underlying role of this gene in protocorm development and differentiation	[78]
IAA载体蛋白基因 IAA carrier protein gene	CtAUX1	无距虾脊兰 C. tsoongiana	qRT-PCR、激素含量检测 qRT-PCR, detection of endogenous hormone content	参与原球茎叶片的起始和发育 Play a key role in the initiation and development of protocorm leaves	[78]
KNOX转录因子 KNOX transcription factor	CtSTM	无距虾脊兰 C. tsoongiana	qRT-PCR	参与原球茎中SAM形成 Involve in shoot apical meristem formation in protocorm	[78]
NAC转录因子 NAC transcription factor	CtCUC2	无距虾脊兰 C. tsoongiana	转录组差异分析 Differential expression gene in transcriptome	参与原球茎生长发育 Involve in the growth and development of protocorm	[78]
MYB转录因子 MYB transcription factor	CtAS1	无距虾脊兰 C. tsoongiana	转录组差异分析 Differential expression gene in transcriptome	参与原球茎生长发育 Involve in the growth and development of protocorm	[78]
泛素结合酶基因 Ubiquitin-conjugating enzyme gene	DoUBC9	铁皮石斛 Dendrobium officinale	cDNA末端快速扩增技术RACE、生物信息学分析、亚细胞定位预测、qRT-PCR Rapid amplification of cDNA ends, bioinformatics analysis, subcellular localization prediction, qRT-PCR	原球茎形成初期细胞快速分裂与生长 Involve in rapid division and growth of cells in early stage of protocorm	[75]
植物转录因子TIFY家族 Plant transcription factor TIFY gene family	DoIAZ3、5、6、8	铁皮石斛 D. officinale	转录组数据分析、qRT-PCR Transcriptome analysis, qRT-PCR	参与原球茎生长发育 Involve in the growth and development of protocorm	[70]
氨基酸通透酶基因 Amino acid permease gene	DoAAP2	铁皮石斛 D. officinale	RACE、巢式PCR、生物信息学分析、qRT-PCR Rapid amplification of cDNA ends(RACE), Nested PCR, bioinformatics analysis, qRT-PCR	参与原球茎根、茎的发育 Play an important role in the development of root and stem of protocorm	[74]
DELLA家族成员 DELLA gene family	DoDELLA	铁皮石斛 D. officinale	生物信息学分析、qRT-PCR Bioinformatics analysis, qRT-PCR	调控种子萌发和原球茎发育 Regulate seed germination and protocorm development	[79]
生长素响应因子 Auxin response factor	DoARF	铁皮石斛 D. officinale	转录组差异分析、激素信号通路富集分析 Transcriptome differences analysis, hormone signal transduction	参与原球茎发育 Involve in the growth and development of protocorm	[69]
硫氧还蛋白超家族成员 Member of thioredoxin superfamily	DoTrxL2	铁皮石斛 D. officinale	转录组差异分析、功能注释、RACE、巢式PCR、 qRT-PCR Transcriptome differences analysis, gene annotations, RACE, Nested PCR, qRT-PCR	促进原球茎形成 Play an important role in the development of protocorm	[69]
胚胎发育晚期丰富蛋白基因 Late embryogenesis abundant protein gene	DoLEA45、DoLEA52	铁皮石斛 D. officinale	基因家族鉴定、蛋白理化性质分析、qRT-PCR Gene family identification, protein physicoch- emical properties analysis, qRT-PCR	原球茎形态建成中可能发挥重要作用 Play an important role in the morphogenesis of protocorm	[73]
泛素结合酶基因 Ubiquitin- conjugating enzyme gene	DoUBC24	铁皮石斛 D. officinale	RACE、亚细胞定位预测、qRT-PCR、进化分析 RACE, subcellular localization prediction, qRT-PCR、phylogenetic analysis	对原球茎不同发育时期具有重要的调控作用 Play an important role in the development of protocorm	[72]
WOX转录因子 WOX transcription factor	DoWOX(1、2、3、4、6、 8、9、10、11)	铁皮石斛 D. officinale	同源比对、顺式作用元件分析、qRT-PCR Homologous comparison, upstream promoter element, qRT-PCR	参与原球茎建成、生长与发育 Play an important role in protocorm development	[84]
体胚发生类受体激酶基因 Somatic embryogenesis receptor-like kinase gene	DoSERK1	铁皮石斛 D. officinale	同源比对、qRT-PCR Homologous comparison, qRT-PCR	参与原球茎SAM和叶片形成 Play an important role in the formation of apical meristem and leaf	[84]
体胚发生类受体激酶基因 Somatic embryogenesis receptor-like kinase gene	DoSERK2	铁皮石斛 D. officinale	同源比对、qRT-PCR Homologous comparison, qRT-PCR	参与原球茎发育 Play an important role in the protocorm development	[84]
胚胎发育相关蛋白基因 Embryogenesis-associated protein gene	DoEMB8	铁皮石斛 D. officinale	RACE、巢式PCR、进化分析、亚细胞定位预测、 qRT-PCR RACE, Nested PCR, phylogenetic analysis, subcellular localization prediction, qRT-PCR	参与种胚生长发育 Play an important role in plant embryonic development	[83]
促分裂原蛋白激酶基因 Mitogen-activated protein kinases gene	DoMPK4	铁皮石斛 D. officinale	RACE、qRT-PCR RACE, qRT-PCR	参与原球茎生长发育 Play an important role in the protocorm development	[71]
富含羟脯氨酸糖蛋白 Hydroxyproline-rich glycoproteins	HRGPs	铁皮石斛 D. officinale	qRT-PCR、免疫荧光标记、免疫印记分析 qRT-PCR, Immunofluorescence staining, immunodot blots	参与共生萌发 Involve in symbiotic germination	[82]
促分裂原蛋白激酶基因 Mitogen-activated protein kinase gene	DoMAPK5	铁皮石斛 D. officinale	qRT-PCR	参与接菌共生种子萌发 Involve in symbiotic seed germination	[81]
ABC转运蛋白 ABC transporter protein	ABCG-PDR、ABCB11	铁皮石斛 D. officinale	基因家族鉴定、进化分析、亚细胞定位、qRT-PCR Gene family identification, phylogenetic analysis, subcellular localization prediction, qRT-PCR	参与激素运输及菌共生种子萌发 Involve in acid and auxin transport	[77]
赤霉素3-氧化酶基因 Gibberellin 3-oxidases gene	DoGA3ox	铁皮石斛 D. officinale	转录组差异分析、序列比对与进化分析、 RACE、qRT-PCR Transcriptome differences analysis, sequence align- ment and Phylogenetic analysis, RACE, qRT-PCR	调控种子萌发 Play a crucial role in the regulation of seed germination	[80]
类原球茎PLB
乙烯响应因子 Ethylene response factor	DoERF5	铁皮石斛 D. officinale	转录组、qRT-PCR、转拟南芥、酵母单杂 Transcriptome analysis, qRT-PCR, transformation of Arabidopsis thaliana, yeast one-hybrid	促类原球茎再生 Enhance the PLB regeneration from PLB and stem explants	[86]
KNOX转录因子 KNOX transcription factor	DoSTM	铁皮石斛 D. officinale	转录组、qRT-PCR、转拟南芥、酵母单杂、转石斛PLB Transcriptome analysis, qRT-PCR, Transformation of Arabidopsis thaliana, yeast one-hybrid, transformation PLB of D. officinale	促类原球茎再生 Enhance the PLB regeneration from PLB and stem explants	[86]
体胚发生类受体激酶基因 Somatic embryogenesis receptor-like kinase genes	PhSERK(1-5)	蝴蝶兰 Phalaenopsis	qRT-PCR	参与PLB诱导和PLB生长发育 Play an important role in PLB induction and subsequent development	[19]
KNOX转录因子 KNOX transcription factor	PaSTM	蝴蝶兰 Phalaenopsis aphrodite	qRT-PCR、同源与异源转化 qRT-PCR, homologous and heterologous transformations	参与PLB的发育 Be important for PLB development	[21]
WOX转录因子 WOX transcription factors	DoWOX(1、2、3、4、9)	铁皮石斛 D. officinale	生物信息学分析、qRT-PCR Bioinformatics analysis, qRT-PCR	潜在的类原球茎发育调控因子 Play an important role in maintenance of PLB state	[90]
AP2转录因子 AP2 transcription factors	DoAP2(2、5、7、8、12)	铁皮石斛 D. officinale	生物信息学分析、qRT-PCR Bioinformatics analysis, qRT-PCR	类原球茎再生 PLB regeneration	[91]

分类 Classification	基因名称 Gene name	物种 Species	鉴定方法 Verification method	功能 Function	参考文献 Reference
原球茎Protocorm
胚胎发育相关基因 Embryonic development related genes	PsSERKs、PsBBMs	白旗兜兰 Paphiopedilum spicerianum	转录组、实时荧光定量PCR(qRT-PCR) Transcriptome analysis、Real-Time PCR(qRT-PCR)	参与原球茎的发育 Involve in the development of protocorm	[76]
SAM形成相关基因 Shoot apical meristem related genes	PsWOX8、PsCLAVATA2、 PsCUC2、PsSCR	白旗兜兰 P. spicerianum	转录组、qRT-PCR Transcriptome analysis、qRT-PCR	参与原球茎中顶端分生组织SAM形成 Involve in shoot apical meristem formation in protocorm	[76]
ABA生物合成基因 ABA biosynthesis genes	CtNCED1、CtZEP	无距虾脊兰 Calanthe tsoongiana	qRT-PCR、激素含量检测 qRT-PCR, detection of endogenous hormone content	参与原球茎发育 Involve in the development of protocorm	[78]
IAA合成关键酶基因 IAA synthesis key enzymes gene	CtTSA1	无距虾脊兰 C. tsoongiana	qRT-PCR、激素含量检测 qRT-PCR, detection of endogenous hormone content	在原球茎发育和分化中具有潜在作用 An underlying role of this gene in protocorm development and differentiation	[78]
IAA载体蛋白基因 IAA carrier protein gene	CtAUX1	无距虾脊兰 C. tsoongiana	qRT-PCR、激素含量检测 qRT-PCR, detection of endogenous hormone content	参与原球茎叶片的起始和发育 Play a key role in the initiation and development of protocorm leaves	[78]
KNOX转录因子 KNOX transcription factor	CtSTM	无距虾脊兰 C. tsoongiana	qRT-PCR	参与原球茎中SAM形成 Involve in shoot apical meristem formation in protocorm	[78]
NAC转录因子 NAC transcription factor	CtCUC2	无距虾脊兰 C. tsoongiana	转录组差异分析 Differential expression gene in transcriptome	参与原球茎生长发育 Involve in the growth and development of protocorm	[78]
MYB转录因子 MYB transcription factor	CtAS1	无距虾脊兰 C. tsoongiana	转录组差异分析 Differential expression gene in transcriptome	参与原球茎生长发育 Involve in the growth and development of protocorm	[78]
泛素结合酶基因 Ubiquitin-conjugating enzyme gene	DoUBC9	铁皮石斛 Dendrobium officinale	cDNA末端快速扩增技术RACE、生物信息学分析、亚细胞定位预测、qRT-PCR Rapid amplification of cDNA ends, bioinformatics analysis, subcellular localization prediction, qRT-PCR	原球茎形成初期细胞快速分裂与生长 Involve in rapid division and growth of cells in early stage of protocorm	[75]
植物转录因子TIFY家族 Plant transcription factor TIFY gene family	DoIAZ3、5、6、8	铁皮石斛 D. officinale	转录组数据分析、qRT-PCR Transcriptome analysis, qRT-PCR	参与原球茎生长发育 Involve in the growth and development of protocorm	[70]
氨基酸通透酶基因 Amino acid permease gene	DoAAP2	铁皮石斛 D. officinale	RACE、巢式PCR、生物信息学分析、qRT-PCR Rapid amplification of cDNA ends(RACE), Nested PCR, bioinformatics analysis, qRT-PCR	参与原球茎根、茎的发育 Play an important role in the development of root and stem of protocorm	[74]
DELLA家族成员 DELLA gene family	DoDELLA	铁皮石斛 D. officinale	生物信息学分析、qRT-PCR Bioinformatics analysis, qRT-PCR	调控种子萌发和原球茎发育 Regulate seed germination and protocorm development	[79]
生长素响应因子 Auxin response factor	DoARF	铁皮石斛 D. officinale	转录组差异分析、激素信号通路富集分析 Transcriptome differences analysis, hormone signal transduction	参与原球茎发育 Involve in the growth and development of protocorm	[69]
硫氧还蛋白超家族成员 Member of thioredoxin superfamily	DoTrxL2	铁皮石斛 D. officinale	转录组差异分析、功能注释、RACE、巢式PCR、 qRT-PCR Transcriptome differences analysis, gene annotations, RACE, Nested PCR, qRT-PCR	促进原球茎形成 Play an important role in the development of protocorm	[69]
胚胎发育晚期丰富蛋白基因 Late embryogenesis abundant protein gene	DoLEA45、DoLEA52	铁皮石斛 D. officinale	基因家族鉴定、蛋白理化性质分析、qRT-PCR Gene family identification, protein physicoch- emical properties analysis, qRT-PCR	原球茎形态建成中可能发挥重要作用 Play an important role in the morphogenesis of protocorm	[73]
泛素结合酶基因 Ubiquitin- conjugating enzyme gene	DoUBC24	铁皮石斛 D. officinale	RACE、亚细胞定位预测、qRT-PCR、进化分析 RACE, subcellular localization prediction, qRT-PCR、phylogenetic analysis	对原球茎不同发育时期具有重要的调控作用 Play an important role in the development of protocorm	[72]
WOX转录因子 WOX transcription factor	DoWOX(1、2、3、4、6、 8、9、10、11)	铁皮石斛 D. officinale	同源比对、顺式作用元件分析、qRT-PCR Homologous comparison, upstream promoter element, qRT-PCR	参与原球茎建成、生长与发育 Play an important role in protocorm development	[84]
体胚发生类受体激酶基因 Somatic embryogenesis receptor-like kinase gene	DoSERK1	铁皮石斛 D. officinale	同源比对、qRT-PCR Homologous comparison, qRT-PCR	参与原球茎SAM和叶片形成 Play an important role in the formation of apical meristem and leaf	[84]
体胚发生类受体激酶基因 Somatic embryogenesis receptor-like kinase gene	DoSERK2	铁皮石斛 D. officinale	同源比对、qRT-PCR Homologous comparison, qRT-PCR	参与原球茎发育 Play an important role in the protocorm development	[84]
胚胎发育相关蛋白基因 Embryogenesis-associated protein gene	DoEMB8	铁皮石斛 D. officinale	RACE、巢式PCR、进化分析、亚细胞定位预测、 qRT-PCR RACE, Nested PCR, phylogenetic analysis, subcellular localization prediction, qRT-PCR	参与种胚生长发育 Play an important role in plant embryonic development	[83]
促分裂原蛋白激酶基因 Mitogen-activated protein kinases gene	DoMPK4	铁皮石斛 D. officinale	RACE、qRT-PCR RACE, qRT-PCR	参与原球茎生长发育 Play an important role in the protocorm development	[71]
富含羟脯氨酸糖蛋白 Hydroxyproline-rich glycoproteins	HRGPs	铁皮石斛 D. officinale	qRT-PCR、免疫荧光标记、免疫印记分析 qRT-PCR, Immunofluorescence staining, immunodot blots	参与共生萌发 Involve in symbiotic germination	[82]
促分裂原蛋白激酶基因 Mitogen-activated protein kinase gene	DoMAPK5	铁皮石斛 D. officinale	qRT-PCR	参与接菌共生种子萌发 Involve in symbiotic seed germination	[81]
ABC转运蛋白 ABC transporter protein	ABCG-PDR、ABCB11	铁皮石斛 D. officinale	基因家族鉴定、进化分析、亚细胞定位、qRT-PCR Gene family identification, phylogenetic analysis, subcellular localization prediction, qRT-PCR	参与激素运输及菌共生种子萌发 Involve in acid and auxin transport	[77]
赤霉素3-氧化酶基因 Gibberellin 3-oxidases gene	DoGA3ox	铁皮石斛 D. officinale	转录组差异分析、序列比对与进化分析、 RACE、qRT-PCR Transcriptome differences analysis, sequence align- ment and Phylogenetic analysis, RACE, qRT-PCR	调控种子萌发 Play a crucial role in the regulation of seed germination	[80]
类原球茎PLB
乙烯响应因子 Ethylene response factor	DoERF5	铁皮石斛 D. officinale	转录组、qRT-PCR、转拟南芥、酵母单杂 Transcriptome analysis, qRT-PCR, transformation of Arabidopsis thaliana, yeast one-hybrid	促类原球茎再生 Enhance the PLB regeneration from PLB and stem explants	[86]
KNOX转录因子 KNOX transcription factor	DoSTM	铁皮石斛 D. officinale	转录组、qRT-PCR、转拟南芥、酵母单杂、转石斛PLB Transcriptome analysis, qRT-PCR, Transformation of Arabidopsis thaliana, yeast one-hybrid, transformation PLB of D. officinale	促类原球茎再生 Enhance the PLB regeneration from PLB and stem explants	[86]
体胚发生类受体激酶基因 Somatic embryogenesis receptor-like kinase genes	PhSERK(1-5)	蝴蝶兰 Phalaenopsis	qRT-PCR	参与PLB诱导和PLB生长发育 Play an important role in PLB induction and subsequent development	[19]
KNOX转录因子 KNOX transcription factor	PaSTM	蝴蝶兰 Phalaenopsis aphrodite	qRT-PCR、同源与异源转化 qRT-PCR, homologous and heterologous transformations	参与PLB的发育 Be important for PLB development	[21]
WOX转录因子 WOX transcription factors	DoWOX(1、2、3、4、9)	铁皮石斛 D. officinale	生物信息学分析、qRT-PCR Bioinformatics analysis, qRT-PCR	潜在的类原球茎发育调控因子 Play an important role in maintenance of PLB state	[90]
AP2转录因子 AP2 transcription factors	DoAP2(2、5、7、8、12)	铁皮石斛 D. officinale	生物信息学分析、qRT-PCR Bioinformatics analysis, qRT-PCR	类原球茎再生 PLB regeneration	[91]

兰科植物原球茎和类原球茎研究进展

Research progress of protocorm and protocorm-like body in Orchidaceae plants

RichHTML

PDF (PC)

可视化

摘要/Abstract

引用本文

使用本文

图/表 2

参考文献 120

相关文章 15

编辑推荐

Metrics

本文评价

[1]	胡心柔, 王梅, 张雅芬, 蔡为明, 金群力. 非生物胁迫对灵芝生长发育及其响应机制的影响[J]. 浙江农业学报, 2025, 37(5): 1182-1190.
[2]	闵江艳, 唐卓磊, 杨雪, 黄小燕, 黄凯丰, 何佩云. 不同干旱-复水模式对苦荞生长与产量的影响[J]. 浙江农业学报, 2024, 36(9): 2000-2009.
[3]	高国际, 龙玲, 宋晓云, 李彦彤, 刘高强, 丁功涛. 亮斑扁角水虻幼虫代替豆粕对北京鸭生长发育和血清生化指标的影响[J]. 浙江农业学报, 2024, 36(8): 1764-1772.
[4]	俞瑞鲜, 胡秀卿, 柳新菊, 汤涛, 吴珉, 吴声敢, 赵学平. 亚致死剂量氰戊菊酯对家蚕生长发育的影响[J]. 浙江农业学报, 2024, 36(2): 264-271.
[5]	叶涛, 孙钦玉, 陈伟立, 单文书, 连文旭, 牛婷婷, 张家侠. 植物病原真菌bZIP基因家族研究进展[J]. 浙江农业学报, 2024, 36(12): 2885-2894.
[6]	廖雪环, 张坷塬, 阿尔力色, 周林, 杨尔倮, 邓俊, 张荣萍. 柯杈肥与功能肥复配对杂交稻生长与产量的影响[J]. 浙江农业学报, 2024, 36(11): 2447-2455.
[7]	尤翠翠, 贺一哲, 徐鹏, 黄亚茹, 王辉, 何海兵, 柯健, 武立权. 高温胁迫对水稻生长发育的伤害效应及其防御对策[J]. 浙江农业学报, 2023, 35(1): 10-22.
[8]	李春梅, 万小荣, 关子盈, 赖晓凤, 罗凯晴, 刘凯. 长链非编码RNA调控植物生长发育与逆境胁迫响应研究进展[J]. 浙江农业学报, 2022, 34(9): 2066-2076.
[9]	杨超, 刘敏竹, 李强, 韩涛, 彭良志, 凌丽俐, 付行政, 淳长品, 曹立, 何义仲. 发光二极管(LED)光质对金秋砂糖橘幼苗生长发育和光合特性的影响[J]. 浙江农业学报, 2022, 34(1): 89-97.
[10]	许娜, 王大海, 杜传印, 杜沙沙, 王晓萌, 张彦, 张玉琴, 吴元华, 管恩森, 石屹. 株距对烟苗生长发育的影响[J]. 浙江农业学报, 2020, 32(8): 1342-1350.
[11]	耿艳飞, 吕明芳. 植物富含半胱氨酸类受体激酶家族研究进展[J]. 浙江农业学报, 2020, 32(12): 2303-2312.
[12]	巫伟峰, 陈孝丑, 陈发兴, 陈春, 张毅智. 基于ITS2序列探讨兰属的DNA条形码鉴定和系统发育关系[J]. 浙江农业学报, 2019, 31(8): 1295-1304.
[13]	刘慧洁, 徐恒, 邱文怡, 李晓芳, 张华, 朱英, 李春寿, 王良超. 转录因子在植物生长发育及非生物逆境响应的作用[J]. 浙江农业学报, 2019, 31(7): 1205-1214.
[14]	程华, 卢凤君, 刘晴. 畜禽种质资源开发利用的全链闭环模式——来自峪口禽业的例证[J]. 浙江农业学报, 2019, 31(10): 1734-1744.
[15]	谢宇凯, 郑许松, 田俊策, 张大羽, 吕仲贤. 不同生存基质对中华淡翅盲蝽生长发育和繁殖的影响[J]. 浙江农业学报, 2018, 30(3): 432-436.