花序结构调控研究进展

doi:10.16420/j.issn.0513-353x.2024-0775

摘要/Abstract

摘要：

植物株型和花序结构受分生组织、开花基因、激素等多种因素的影响。本文综述了影响拟南芥、水稻、番茄等花序结构发育进程的调控网络机制，发现在边界区特异性表达的基因是花序分生组织（inflorescence meristem，IM）和花分生组织（flower meristem，FM）正常发育的基础；IM的发育和FM的确定都需要TFL1、LFY、AP1、SOC1、AGL24、SVP、SEP4等关键调节因子的相互作用；WUS-CLV和WUS-AG反馈系统通过影响干细胞的活性来确定花序结构和花朵数量；而IAA、GA和CTK等激素是调控网络中的关键环节；本文还对茄科蔬菜花序复杂性状形成机理进行了探讨，对未来的研究进行了展望，以期对花序形态建成等方面的研究和作物育种提供借鉴。

关键词: 花序发育, 花芽分化, 开花调控, 植物激素

Abstract:

Meristem，flowering genes，hormones，and other factors influence plant type and inflorescence structure. This paper reviews the regulatory network mechanisms affecting the development of inflorescence structure in Arabidopsis thaliana，rice，and tomato. It is found that the genes specifically expressed in the boundary area are the basis for the normal development of inflorescence meristem（IM）and flower meristem（FM）. The development of IM and the determination of FM require the interaction of key regulatory factors such as TFL1，LFY，AP1，SOC1，AGL24，SVP and SEP4. WUS-CLV and WUS-AG feedback systems determine inflorescence structure and flower number by influencing the activity of stem cells. Hormones such as IAA，GA and CTK are the key links in these regulatory networks. In this paper，the formation mechanism of complex inflorescence characters of Solanaceae vegetables was also discussed，and future research is expected to provide a reference for the research on inflorescence morphogenesis and crop breeding.

Key words: inflorescence development, flower bud differentiation, flowering regulation, plant hormone

李美玉, 王本启, 黄树苹, 陈霞, 谈杰, 张洪源, 王俊良, 陈蓉, 张俊红, 张敏. 花序结构调控研究进展[J]. 园艺学报, 2025, 52(8): 2249-2269.

LI Meiyu, WANG Benqi, HUANG Shuping, CHEN Xia, TAN Jie, ZHANG Hongyuan, WANG Junliang, CHEN Rong, ZHANG Junhong, and ZHANG Min. The Research Progress of Inflorescence Structure Regulation[J]. Acta Horticulturae Sinica, 2025, 52(8): 2249-2269.

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参考文献 169

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基因名Gene name	基因ID Gene ID	基因类型Gene type	功能描述Fnctional description	参考文献Rference
CUC1	AT3G15170	NAC（无顶端分生组织）结构域转录调节因子超家族蛋白 Transcriptional regulator superfamily protein of NAC （apical meristem-free）domain	顶端分生组织、STM形成所必需 It is necessary for the formation of apical meristem and STM	Ishida et al.,2000
CUC2	AT5G53950		调节腋生分生组织起始 Regulation of axillary meristem initiation	Aida et al.,1997
CUC3	AT1G76420		调控边界区形成 Regulating the formation of boundary area	Raman et al.,2008
miR164A	AT2G47585	miRNA初级转录本 MiRNA primary transcript	编码靶向包含NAC结构域的基因 Coding targets genes containing NAC domain.	Allen et al.,2005
MiR164B	AT5G01747
MiR164C	AT5G27807
miR156			负调控IM起始 Negative regulation of IM initiation	Wang et al.,2015
TFL1	AT5G03840	PEBP家族蛋白 PEBP family protein	维持IM特性，促进分枝 Maintain IM characteristics and promote branching	Shannon & Meeks-Wagner,1991
SP	Solyc06g074350		促进分枝，维持茎尖特性 Promote branching and maintain shoot tip characteristics.	Pnueli et al.,1998
FT	AT1G65480		促进开花Promote flowering	Kardailsky et al.,1999
CAL	AT1G26310	MADS-box转录因子家族 MADS-box transcription factor family	促进FM的形成 Promote the formation of FM	Elena et al.,2006
AGAMOUS	LOC110785792		负调控干细胞的积累 Negative regulation of stem cell accumulation	Sun et al.,2009
SVP	AT2G22540		抑制开花Suppress flowering	Hartmann et al.,2000
FUL	AT5G60910		促进开花Promote flowering	Yamaguchi et al.,2009
FLC	AT5G10140		促进开花Promote flowering	Buzas et al.,2021
SEP3	AT1G24260		促进FM转换Promote FM conversion	Hugouvieux et al.,2018
SEP4	AT2G03710		确定FM和花器官的形成 Determine FM and floral organ formation	Gary et al.,2004
TAW1	LOC_Os10g33780	ALOG家族蛋白 ALOG family protein	使花序不确定性生长 Make inflorescence grow indefinitely.	Yoshida et al.,2013
TMF	Solyc09g090180	ALOG家族蛋白 ALOG family protein	番茄花序的复杂度 Complexity of tomato inflorescence	Macalister et al.,2012
LFY	AT5G61850	LEAFY家族 LEAFY family	促进FM和花的形成 Promote FM and flower formation	Yamaguchi et al.,2016
DA1	AT1G19270	含LIM结构域的蛋白 LIM domain-containing protein	调控腋生分生组织的起始 Regulating the initiation of axillary meristem	Koch et al.,2006
UBP15	AT1G17110	泛素特异性蛋白酶15 Ubiquitin-specific protease 15	抑制腋生分行组织的启动 Inhibit the initiation of axillary branch tissue	Yan et al., 2000
DPA4/NGAL3	AT5G06250	AP2/B3样转录因子家族蛋白 AP2/B3-like ranscription factor family protein	花序结构调节转录阻遏物 Inflorescence structure regulates transcription repressor	Riechmann et al.,2000
SOD7/NGAL2	AT3G11580		腋生分生组织形成所必需 Necessary for axillary meristem formation
LAS	AT1G55580	GRAS家族转录因子 GRAS family transcription factors	调节腋生分生组织的起始 Regulation of axillary meristem initiation	Greb et al.,2003
MOC1	LOC127776671	蛋白质MONOCULM 1 Protein MONOCULM 1	正调控水稻分蘖 Positive regulation of rice tillering	Liao et al.,2019
FON1	LOC_Os06g50340	富含亮氨酸的重复受体样激酶蛋白花器官 Leucine-rich repetitive receptor-like kinase protein flower organ	控制花分生组织大小 Controlling the size of flower meristem	Moon et al.,2006
OSH1	LOC_Os03g51690	同源盒蛋白 Knotted-1-like 6 Homocassette protein Knotted-1-like 6	诱导产生腋下小花Induced axillary floret	Zhang et al.,2017
LAX1	LOC_Os01g61480	转录因子LAX PANICLE 1样 Transcription factor LAX PANICLE 1-like	正向调控穗数 Forward regulation of paniCLE number	Matin & Kang,2012
LS	Solyc07g066250	VHIID蛋白 VHIID protein	调控番茄分枝数 Regulating the number of tomato branches	Schumacher et al.,1999
BOP2	AT2G41370	锚蛋白重复家族蛋白/含BTB/POZ结构域的蛋白 Ankyrin repeat family protein/protein containing BTB/POZ domain	促进FM的形成 Promote the formation of FM	Scofield et al.,2018
WUS	AT2G17950	同源结构域样超家族蛋白 Homologous domain-like superfamily protein	激活、维持干细胞表达 Activate and maintain the expression of stem cells	Laux et al.,1996
CLV3	AT2G27250	CLE基因家族 CLE gene family	负调节干细胞的积累 Negative regulation of stem cell accumulation	Clark et al.,1996
CLV1	AT1G75820	富含亮氨酸受体样蛋白激酶家族蛋白 Leucine-rich receptor-like protein kinase family protein	促进FM的形成 Promote the formation of FM	Clark et al.,1997
RCN1	LOC_Os11g05470	aBC转运蛋白G家族成员5 ABC transporter G family member 5	负向调控花发育 Negative regulation of flower development	Hanzawa et al.,2005
SFT	Solyc03g063100	CETS家族蛋白 CETS family protein	促进番茄开花Promote tomato flowering	Lifschitz et al.,2014
PAP2	LOC_Os03g54170		正调控小穗分生组织的形成 Positive regulation of spikelet meristem formation	Liu et al.,2013
CO	AT5G15840	B-box型锌指蛋白 B-box zinc finger protein	长日照下促进开花 Promote flowering under long sunshine	Jung et al.,2012
AP2	AT4G36920	整合酶型DNA结合超家族蛋白 Integrase-type DNA-binding superfamily proteins	确定FM和花器官的形成 Determine FM and floral organ formation	Huang et al.,2017
SPL	AT4G27330	无孢子细胞SPL Sporeless cell SPL	促进FM转换Promote FM conversion	Bencivenga et al.,2012
KNU	AT5G14010	C2H2型锌指蛋白 C2H2 zinc finger protein	在FM中介导WUS抑制 WUS inhibition mediated by FM	Sun et al.,2019
UFO	AT1G30950	F-box家族蛋白 F-box family protein	促进FM的转换和顶花的形成 Promote FM conversion and the formation of top flower	Hepworth et al.,2006
BOP1	Solyc04g040220	BTB-ankryin转录因子 BTB-ankryin transcription factor	促进分生组织成熟 Promote meristem maturation	Xu et al.,2016
BOP2	Solyc10g079460
BOP3	Solyc10g079750

基因名Gene name	基因ID Gene ID	基因类型Gene type	功能描述Fnctional description	参考文献Rference
CUC1	AT3G15170	NAC（无顶端分生组织）结构域转录调节因子超家族蛋白 Transcriptional regulator superfamily protein of NAC （apical meristem-free）domain	顶端分生组织、STM形成所必需 It is necessary for the formation of apical meristem and STM	Ishida et al.,2000
CUC2	AT5G53950		调节腋生分生组织起始 Regulation of axillary meristem initiation	Aida et al.,1997
CUC3	AT1G76420		调控边界区形成 Regulating the formation of boundary area	Raman et al.,2008
miR164A	AT2G47585	miRNA初级转录本 MiRNA primary transcript	编码靶向包含NAC结构域的基因 Coding targets genes containing NAC domain.	Allen et al.,2005
MiR164B	AT5G01747
MiR164C	AT5G27807
miR156			负调控IM起始 Negative regulation of IM initiation	Wang et al.,2015
TFL1	AT5G03840	PEBP家族蛋白 PEBP family protein	维持IM特性，促进分枝 Maintain IM characteristics and promote branching	Shannon & Meeks-Wagner,1991
SP	Solyc06g074350		促进分枝，维持茎尖特性 Promote branching and maintain shoot tip characteristics.	Pnueli et al.,1998
FT	AT1G65480		促进开花Promote flowering	Kardailsky et al.,1999
CAL	AT1G26310	MADS-box转录因子家族 MADS-box transcription factor family	促进FM的形成 Promote the formation of FM	Elena et al.,2006
AGAMOUS	LOC110785792		负调控干细胞的积累 Negative regulation of stem cell accumulation	Sun et al.,2009
SVP	AT2G22540		抑制开花Suppress flowering	Hartmann et al.,2000
FUL	AT5G60910		促进开花Promote flowering	Yamaguchi et al.,2009
FLC	AT5G10140		促进开花Promote flowering	Buzas et al.,2021
SEP3	AT1G24260		促进FM转换Promote FM conversion	Hugouvieux et al.,2018
SEP4	AT2G03710		确定FM和花器官的形成 Determine FM and floral organ formation	Gary et al.,2004
TAW1	LOC_Os10g33780	ALOG家族蛋白 ALOG family protein	使花序不确定性生长 Make inflorescence grow indefinitely.	Yoshida et al.,2013
TMF	Solyc09g090180	ALOG家族蛋白 ALOG family protein	番茄花序的复杂度 Complexity of tomato inflorescence	Macalister et al.,2012
LFY	AT5G61850	LEAFY家族 LEAFY family	促进FM和花的形成 Promote FM and flower formation	Yamaguchi et al.,2016
DA1	AT1G19270	含LIM结构域的蛋白 LIM domain-containing protein	调控腋生分生组织的起始 Regulating the initiation of axillary meristem	Koch et al.,2006
UBP15	AT1G17110	泛素特异性蛋白酶15 Ubiquitin-specific protease 15	抑制腋生分行组织的启动 Inhibit the initiation of axillary branch tissue	Yan et al., 2000
DPA4/NGAL3	AT5G06250	AP2/B3样转录因子家族蛋白 AP2/B3-like ranscription factor family protein	花序结构调节转录阻遏物 Inflorescence structure regulates transcription repressor	Riechmann et al.,2000
SOD7/NGAL2	AT3G11580		腋生分生组织形成所必需 Necessary for axillary meristem formation
LAS	AT1G55580	GRAS家族转录因子 GRAS family transcription factors	调节腋生分生组织的起始 Regulation of axillary meristem initiation	Greb et al.,2003
MOC1	LOC127776671	蛋白质MONOCULM 1 Protein MONOCULM 1	正调控水稻分蘖 Positive regulation of rice tillering	Liao et al.,2019
FON1	LOC_Os06g50340	富含亮氨酸的重复受体样激酶蛋白花器官 Leucine-rich repetitive receptor-like kinase protein flower organ	控制花分生组织大小 Controlling the size of flower meristem	Moon et al.,2006
OSH1	LOC_Os03g51690	同源盒蛋白 Knotted-1-like 6 Homocassette protein Knotted-1-like 6	诱导产生腋下小花Induced axillary floret	Zhang et al.,2017
LAX1	LOC_Os01g61480	转录因子LAX PANICLE 1样 Transcription factor LAX PANICLE 1-like	正向调控穗数 Forward regulation of paniCLE number	Matin & Kang,2012
LS	Solyc07g066250	VHIID蛋白 VHIID protein	调控番茄分枝数 Regulating the number of tomato branches	Schumacher et al.,1999
BOP2	AT2G41370	锚蛋白重复家族蛋白/含BTB/POZ结构域的蛋白 Ankyrin repeat family protein/protein containing BTB/POZ domain	促进FM的形成 Promote the formation of FM	Scofield et al.,2018
WUS	AT2G17950	同源结构域样超家族蛋白 Homologous domain-like superfamily protein	激活、维持干细胞表达 Activate and maintain the expression of stem cells	Laux et al.,1996
CLV3	AT2G27250	CLE基因家族 CLE gene family	负调节干细胞的积累 Negative regulation of stem cell accumulation	Clark et al.,1996
CLV1	AT1G75820	富含亮氨酸受体样蛋白激酶家族蛋白 Leucine-rich receptor-like protein kinase family protein	促进FM的形成 Promote the formation of FM	Clark et al.,1997
RCN1	LOC_Os11g05470	aBC转运蛋白G家族成员5 ABC transporter G family member 5	负向调控花发育 Negative regulation of flower development	Hanzawa et al.,2005
SFT	Solyc03g063100	CETS家族蛋白 CETS family protein	促进番茄开花Promote tomato flowering	Lifschitz et al.,2014
PAP2	LOC_Os03g54170		正调控小穗分生组织的形成 Positive regulation of spikelet meristem formation	Liu et al.,2013
CO	AT5G15840	B-box型锌指蛋白 B-box zinc finger protein	长日照下促进开花 Promote flowering under long sunshine	Jung et al.,2012
AP2	AT4G36920	整合酶型DNA结合超家族蛋白 Integrase-type DNA-binding superfamily proteins	确定FM和花器官的形成 Determine FM and floral organ formation	Huang et al.,2017
SPL	AT4G27330	无孢子细胞SPL Sporeless cell SPL	促进FM转换Promote FM conversion	Bencivenga et al.,2012
KNU	AT5G14010	C2H2型锌指蛋白 C2H2 zinc finger protein	在FM中介导WUS抑制 WUS inhibition mediated by FM	Sun et al.,2019
UFO	AT1G30950	F-box家族蛋白 F-box family protein	促进FM的转换和顶花的形成 Promote FM conversion and the formation of top flower	Hepworth et al.,2006
BOP1	Solyc04g040220	BTB-ankryin转录因子 BTB-ankryin transcription factor	促进分生组织成熟 Promote meristem maturation	Xu et al.,2016
BOP2	Solyc10g079460
BOP3	Solyc10g079750