植物维管形成层发育及其调控的研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2022-0865

摘要/Abstract

摘要：

维管形成层是植物的次生分生组织，它的活动促进植物侧向生长。近年来，大量研究成果加深了对维管形成层的了解，但与顶端分生组织相比，对维管形成层还是知之甚少。遗传与分子生物学研究发现，形成层的增殖与分化受多因素调控，包括长距离激素信号、短距离肽信号及两者之间的相互作用。除此之外，各种转录因子和microRNAs在维管形成层活动的调控过程中也发挥着关键作用。本文主要阐述了维管形成层发育及调控其增殖分化的新发现，并且对该领域目前的研究现状以及未来研究的重点方向进行了总结与展望。

关键词: 维管形成层, 长距离激素信号, 短距离肽信号, 转录因子, microRNAs

Abstract:

The vascular cambium, part of the secondary lateral meristem, contributes to the lateral growth of a plant. In recent years, many studies have strengthened our understanding of the vascular cambium. However, we still know far less about the vascular cambium than the apical meristem. Genetic and molecular studies have revealed that the proliferation and differentiation of cambium is regulated by many factors, including long-distance hormone signals, short-range peptide signals, and interactions between them. In addition, a large number of transcription factors and microRNAs also play a crucial role in the regulation of the vascular cambium activities. This review focuses on the novel discoveries on the development of vascular cambium and its regulation of proliferation and differentiation. The review also summarizes the current research and prospects for future research in this field.

Key words: vascular cambium, long-distance hormone signaling, short-range peptide signal, transcription factor, microRNAs

葛颜锐, 赵冉, 徐静, 李若凡, 胡云涛, 李瑞丽. 植物维管形成层发育及其调控的研究进展[J]. 生物技术通报, 2023, 39(3): 13-25.

GE Yan-rui, ZHAO Ran, XU Jing, LI Ruo-fan, HU Yun-tao, LI Rui-li. Advances in the Development and Regulation of Vascular Cambium[J]. Biotechnology Bulletin, 2023, 39(3): 13-25.

图/表 3

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名称 Name	表达部位 Location	受体 Receptor	功能 Function	参考文献 Reference
CLE41/44/TDIF	韧皮部 Phloem	PXY/TDR	促进维管形成层细胞分裂，抑制形成层分化 Promote vascular cambium cell division and inhibit cambium differentiation	[43]
CLE41/44/TDIF	韧皮部 Phloem	XVP		[56]
CLE9/10	木质部前体细胞 Xylem precursor cells	HSL1	抑制气孔细胞的分裂 Inhibit stomatal cell division	[48]
CLE9/10	木质部前体细胞 Xylem precursor cells	BAM	抑制形成层细胞平周分裂 Inhibit periferential division of cambial cells	[48⇓-50]
CLE45	原生韧皮部 Primary phloem	BAM3	负调控拟南芥原韧皮部分化 Negatively regulate of Arabidopsis prophloem differentiation	[51,53]
CLE25	韧皮部 Phloem	CLERK- CLV2	促进韧皮部的启动 Promote phloem initiation	[54]

名称 Name	表达部位 Location	受体 Receptor	功能 Function	参考文献 Reference
CLE41/44/TDIF	韧皮部 Phloem	PXY/TDR	促进维管形成层细胞分裂，抑制形成层分化 Promote vascular cambium cell division and inhibit cambium differentiation	[43]
CLE41/44/TDIF	韧皮部 Phloem	XVP		[56]
CLE9/10	木质部前体细胞 Xylem precursor cells	HSL1	抑制气孔细胞的分裂 Inhibit stomatal cell division	[48]
CLE9/10	木质部前体细胞 Xylem precursor cells	BAM	抑制形成层细胞平周分裂 Inhibit periferential division of cambial cells	[48⇓-50]
CLE45	原生韧皮部 Primary phloem	BAM3	负调控拟南芥原韧皮部分化 Negatively regulate of Arabidopsis prophloem differentiation	[51,53]
CLE25	韧皮部 Phloem	CLERK- CLV2	促进韧皮部的启动 Promote phloem initiation	[54]

名称 Name	表达部位 Location	靶基因 Target gene	功能 Function	参考文献 Reference
MP/ARF5	原形成层 Procambia	WOX4	促进木质部分化 Promote xylem differentiation	[15]
MP/ARF5	原形成层 Procambia	TMO5	控制维管组织的确定，促进原形成层周壁细胞分裂 Control the determination of vascular tissue, and promote the division of periparietal cells in the procambium	[26⇓-28]
TMO5-LHW	初生木质部 Primary xylem	LOG3/4	控制维管组织的确定，促进原形成层周壁细胞分裂 Control the determination of vascular tissue, and promote the division of periparietal cells in the procambium	[29]
TMO5-LHW	初生木质部 Primary xylem	DOF2.1	促进原形成层细胞分裂 Promote the division of the protocambial cells	[30]
WOX4/14	原形成层、维管形成层 Procambia and vascular cambium	TMO6	促进形成层增殖，参与维管束形状的确定 Promote cambium proliferation, and participate in the determination of vascular bundle shape	[59-60]
WOX4/14	原形成层、维管形成层 Procambia and vascular cambium	LBD4
TMO6	维管形成层 Vascular cambium	LBD4
SOC1	顶端分生组织、原形成层 Apical meristem and procambia	AHL15	促进开花，影响分生组织的确定性，抑制次生生长 Promote flowering, affect the certainty of meristem, and inhibit secondary growth	[62-63]
FUL	顶端分生组织、原形成层 Apical meristem and procambia	AHL15		[62-63]
AHL15	维管形成层 Vascular cambium	IPT3/7、LOG5	抑制腋芽分生组织成熟，延长寿命，促进形成层活性，促进次生木质部的发育 Inhibit the maturation of axillary bud meristem, prolong the life span, promote the activity of cambium, and promote the development of secondary xylem	[64]
XVP	维管形成层 Vascular cambium	CLE44	促进木质部分化，维持维管稳定 Promote xylem differentiation, and maintain vascular stability	[65-66]
EPFL4/6	内皮层 Endodermis	ER/ERL1	促进原形成层发育 Promote the development of procambium	[68⇓-70]
GRF	维管形成层、短暂扩充细胞 Vascular cambium、transient amplifying cell	PLT	促进生长 Promote growing	[71]

名称 Name	表达部位 Location	靶基因 Target gene	功能 Function	参考文献 Reference
MP/ARF5	原形成层 Procambia	WOX4	促进木质部分化 Promote xylem differentiation	[15]
MP/ARF5	原形成层 Procambia	TMO5	控制维管组织的确定，促进原形成层周壁细胞分裂 Control the determination of vascular tissue, and promote the division of periparietal cells in the procambium	[26⇓-28]
TMO5-LHW	初生木质部 Primary xylem	LOG3/4	控制维管组织的确定，促进原形成层周壁细胞分裂 Control the determination of vascular tissue, and promote the division of periparietal cells in the procambium	[29]
TMO5-LHW	初生木质部 Primary xylem	DOF2.1	促进原形成层细胞分裂 Promote the division of the protocambial cells	[30]
WOX4/14	原形成层、维管形成层 Procambia and vascular cambium	TMO6	促进形成层增殖，参与维管束形状的确定 Promote cambium proliferation, and participate in the determination of vascular bundle shape	[59-60]
WOX4/14	原形成层、维管形成层 Procambia and vascular cambium	LBD4
TMO6	维管形成层 Vascular cambium	LBD4
SOC1	顶端分生组织、原形成层 Apical meristem and procambia	AHL15	促进开花，影响分生组织的确定性，抑制次生生长 Promote flowering, affect the certainty of meristem, and inhibit secondary growth	[62-63]
FUL	顶端分生组织、原形成层 Apical meristem and procambia	AHL15		[62-63]
AHL15	维管形成层 Vascular cambium	IPT3/7、LOG5	抑制腋芽分生组织成熟，延长寿命，促进形成层活性，促进次生木质部的发育 Inhibit the maturation of axillary bud meristem, prolong the life span, promote the activity of cambium, and promote the development of secondary xylem	[64]
XVP	维管形成层 Vascular cambium	CLE44	促进木质部分化，维持维管稳定 Promote xylem differentiation, and maintain vascular stability	[65-66]
EPFL4/6	内皮层 Endodermis	ER/ERL1	促进原形成层发育 Promote the development of procambium	[68⇓-70]
GRF	维管形成层、短暂扩充细胞 Vascular cambium、transient amplifying cell	PLT	促进生长 Promote growing	[71]