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

doi:10.3969/j.issn.1004-1524.20240581

Acta Agriculturae Zhejiangensis ›› 2025, Vol. 37 ›› Issue (6): 1372-1389.DOI: 10.3969/j.issn.1004-1524.20240581

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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

Abstract

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

CLC Number:

S682.31

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.

Figures/Tables 2

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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]

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

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