Mechanism of peroxisome proliferation in different organisms

doi:10.3969/j.issn.1004-1524.20231108

Acta Agriculturae Zhejiangensis ›› 2024, Vol. 36 ›› Issue (12): 2870-2884.DOI: 10.3969/j.issn.1004-1524.20231108

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Mechanism of peroxisome proliferation in different organisms

HAN Yatao¹(), WANG Mengjing¹(), LU Ziqi¹^,², WANG Jiaoyu²^,^*(), LI Ling¹^,^*()

1. College of Advanced Agricultural Sciences, Zhejiang A&F University, Hangzhou 311300, China
2. Institute of Plant Protection and Microbiology, Zhejiang Academy of Agricultural Sciences, Hangzhou 310021, China

Received:2023-09-15 Online:2024-12-25 Published:2024-12-27

Abstract

Abstract:

Peroxisomes are organelles that are commonly found in eukaryotic cells and are involved in a variety of metabolic processes, especially lipid metabolism. The number and volume of peroxisomes will change in response to changes in the external environment, and their proliferation process is realized mainly through de novo synthesis and proliferation (division) processes. The de novo synthesis process involves the formation of peroxisome precursors (a vesicle-like structure) by the endoplasmic reticulum through budding reproduction, and the recruitment of membrane proteins as well as matrix proteins in the cytoplasm by the precursor vesicles, which eventually become mature peroxisomes. PEX3, PEX16 and PEX19 gene are mainly involved in the de novo production of peroxisomes and play an important role in the biosynthesis of peroxisomal membrane proteins (PMPs). Peroxisome proliferation (division) includes several processes such as peroxisome polarization, membrane protrusion and elongation, matrix protein transport, membrane contraction and division. PEX11 gene encodes a peroxisomal membrane elongation factor that is involved in membrane polarization, protrusion and elongation during peroxisome proliferation. In addition, Vps1 and Dnm1 regulate the morphology, replication and abundance of peroxisomes, and Inp1 and Inp2 are involved in the inheritance of peroxisomes. In this review, we summarize the peroxisome proliferation processes and related gene functions in different organisms to provide ideas for the expectation of peroxisome utilization.

Key words: peroxisome, de novo synthesis, proliferation, PEX gene

CLC Number:

HAN Yatao, WANG Mengjing, LU Ziqi, WANG Jiaoyu, LI Ling. Mechanism of peroxisome proliferation in different organisms[J]. Acta Agriculturae Zhejiangensis, 2024, 36(12): 2870-2884.

Figures/Tables 7

References 109

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物种名称 Species name	基因 Gene	数量 Number	功能 Function
哺乳动物 Mammal	PEX3	1	启动过氧化物酶体前体囊泡的形成,并参与转运剩余PMPs进入过氧化物酶体前体 Initiates the formation of peroxisome precursor vesicles and participates in the transport of residual PMPs into peroxisome prebodies
	PEX16	1	充当PMP受体,帮助协调将这些PMP分类为过氧化物酶体前体,同时是Pex3p插入内质网和过氧化物酶体所必需的 Acts as a PMP receptor, helping to coordinate the classification of these PMPs into peroxisome precursors, and is simultaneously necessary for the insertion of Pex3p into the endoplasmic reticulum and peroxisomes
	PEX19	1	转运剩余PMPs至过氧化物酶体前体 Transports residual PMPs to peroxisome precursors
酿酒酵母 Saccharomyces cerevisiae	PEX3	2	作为PMPs的转运蛋白,参与某些PMPs在前体囊泡的定位 Acts as a transporter protein for PMPs and is involved in the localization of certain PMPs in precursor vesicles
	PEX16	0	—
	PEX19	1	是新合成的Ⅰ类PMPs转运的重要伴侣和受体,参与PMPs的定位 An important chaperone and receptor for the transport of newly synthesized class Ⅰ PMPs, involved in the localization of PMPs
耶式酵母 Yarrowia lipolytica	PEX3	1	作为PMPs的转运蛋白,参与某些PMPs在前体囊泡的定位 Acts as a transporter protein for PMPs and is involved in the localization of certain PMPs in precursor vesicles
	PEX16	1	帮助过氧化物酶体膜蛋白的正确定位以及过氧化物酶的分裂 Aids in the proper localization of peroxisomal membrane proteins and the division of peroxisomal enzymes
	PEX19	1	缺失后仍能形成形态可识别的过氧化物酶体 Formation of morphologically recognizable peroxisomes despite deletions
稻瘟病菌 Magnaporthe oryzae	PEX3	1	与PEX19相互作用,对于PMPs的插入起着至关重要的作用 Interacts with PEX19 and plays a crucial role for the insertion of PMPs
	PEX16	1	是PMPs插入和输入所必需的基因 It’s a gene that is required for PMPs insertion and input
	PEX19	1	保证过氧化物异构膜蛋白和过氧化物异构基质蛋白的正确定位,缺失后会造成脂质代谢和活性氧耐受的缺陷,抑制气生菌丝的生长和孢子形成,降低分生孢子的形态和活力 Ensures the correct localization of peroxisomal membrane proteins and peroxisomal matrix proteins, the absence of which causes defects in lipid metabolism and reactive oxygen tolerance, inhibits aerial mycelium growth and spore formation, and reduces conidial morphology and viability
拟南芥 Arabidopsis thaliana	PEX3	2	被认为是PEX19的一种对接因子 Considered a docking factor for PEX19
	PEX16	1	作为PEX3和其他Ⅱ型PMP受体,作为内质网中PMPs受体,协调这些PMPs分类到过氧化物酶体前体上,同时参与基质蛋白的转运过程 Acts as a receptor for PEX3 and other type Ⅱ PMPs in the endoplasmic reticulum and coordinates the sorting of these PMPs onto peroxisomal precursors, as well as being involved in the transport of matrix proteins
	PEX19	2	是PMP的胞质伴侣,将PMP递送至内质网或过氧化物酶体膜,参与Ⅰ型PMPs的输入 It is a cytoplasmic chaperone for PMPs, delivers PMPs to the endoplasmic reticulum or peroxisomal membrane, and participates in the import of type Ⅰ PMPs

物种名称 Species name	基因 Gene	数量 Number	功能 Function
哺乳动物 Mammal	PEX3	1	启动过氧化物酶体前体囊泡的形成,并参与转运剩余PMPs进入过氧化物酶体前体 Initiates the formation of peroxisome precursor vesicles and participates in the transport of residual PMPs into peroxisome prebodies
	PEX16	1	充当PMP受体,帮助协调将这些PMP分类为过氧化物酶体前体,同时是Pex3p插入内质网和过氧化物酶体所必需的 Acts as a PMP receptor, helping to coordinate the classification of these PMPs into peroxisome precursors, and is simultaneously necessary for the insertion of Pex3p into the endoplasmic reticulum and peroxisomes
	PEX19	1	转运剩余PMPs至过氧化物酶体前体 Transports residual PMPs to peroxisome precursors
酿酒酵母 Saccharomyces cerevisiae	PEX3	2	作为PMPs的转运蛋白,参与某些PMPs在前体囊泡的定位 Acts as a transporter protein for PMPs and is involved in the localization of certain PMPs in precursor vesicles
	PEX16	0	—
	PEX19	1	是新合成的Ⅰ类PMPs转运的重要伴侣和受体,参与PMPs的定位 An important chaperone and receptor for the transport of newly synthesized class Ⅰ PMPs, involved in the localization of PMPs
耶式酵母 Yarrowia lipolytica	PEX3	1	作为PMPs的转运蛋白,参与某些PMPs在前体囊泡的定位 Acts as a transporter protein for PMPs and is involved in the localization of certain PMPs in precursor vesicles
	PEX16	1	帮助过氧化物酶体膜蛋白的正确定位以及过氧化物酶的分裂 Aids in the proper localization of peroxisomal membrane proteins and the division of peroxisomal enzymes
	PEX19	1	缺失后仍能形成形态可识别的过氧化物酶体 Formation of morphologically recognizable peroxisomes despite deletions
稻瘟病菌 Magnaporthe oryzae	PEX3	1	与PEX19相互作用,对于PMPs的插入起着至关重要的作用 Interacts with PEX19 and plays a crucial role for the insertion of PMPs
	PEX16	1	是PMPs插入和输入所必需的基因 It’s a gene that is required for PMPs insertion and input
	PEX19	1	保证过氧化物异构膜蛋白和过氧化物异构基质蛋白的正确定位,缺失后会造成脂质代谢和活性氧耐受的缺陷,抑制气生菌丝的生长和孢子形成,降低分生孢子的形态和活力 Ensures the correct localization of peroxisomal membrane proteins and peroxisomal matrix proteins, the absence of which causes defects in lipid metabolism and reactive oxygen tolerance, inhibits aerial mycelium growth and spore formation, and reduces conidial morphology and viability
拟南芥 Arabidopsis thaliana	PEX3	2	被认为是PEX19的一种对接因子 Considered a docking factor for PEX19
	PEX16	1	作为PEX3和其他Ⅱ型PMP受体,作为内质网中PMPs受体,协调这些PMPs分类到过氧化物酶体前体上,同时参与基质蛋白的转运过程 Acts as a receptor for PEX3 and other type Ⅱ PMPs in the endoplasmic reticulum and coordinates the sorting of these PMPs onto peroxisomal precursors, as well as being involved in the transport of matrix proteins
	PEX19	2	是PMP的胞质伴侣,将PMP递送至内质网或过氧化物酶体膜,参与Ⅰ型PMPs的输入 It is a cytoplasmic chaperone for PMPs, delivers PMPs to the endoplasmic reticulum or peroxisomal membrane, and participates in the import of type Ⅰ PMPs

类别 Form	物种 Specie	蛋白 Protein	功能 Function	参考文献 Reference
过氧化物酶体膜的生物起源 Biological origin of	丝状真菌 Filamentous fungi	Pex3	Pex19膜锚定器 Pex19 membrane anchor	[71,73]
peroxisomal membranes	酵母 Yeast	Pex3	Pex19膜锚定器 Pex19 membrane anchor	[74]
	丝状真菌 Filamentous fungi	Pex19	mPTS受体 mPTS receptor	[71,73]
	酵母 Yeast	Pex19	mPTS受体 mPTS receptor	[74]
基质蛋白导入 Matrix protein introduction	丝状真菌 Filamentous fungi	Pex1,Pex6	参与受体循环的AAA型ATP酶 AAA-type ATPases involved in the receptor cycle	[71,73]
	酵母 Yeast	Pex1,Pex6	参与受体循环的AAA型ATP酶 AAA-type ATPases involved in the receptor cycle	[74]
	丝状真菌 Filamentous fungi	Pex2,Pex10,Pex12	参与受体泛素化(连接酶)的环复合体成分 Components of the ring complex involved in receptor ubiquitination (ligase)	[71,73]
	酵母 Yeast	Pex2,Pex10,Pex12	参与受体泛素化(连接酶)的环复合体成分 Components of the ring complex involved in receptor ubiquitination (ligase)	[74]
	丝状真菌 Filamentous fungi	Pex4	参与受体泛素化的泛素结合酶 Ubiquitin-binding enzymes involved in receptor ubiquitination	[71,73]
	酵母 Yeast	Pex4	参与受体泛素化的泛素结合酶 Ubiquitin-binding enzymes involved in receptor ubiquitination	[74]
	丝状真菌 Filamentous fungi	Pex5,Pex9	PTS1受体 PTS1 receptor	[71,73]
	酵母 Yeast	Pex5	PTS1受体 PTS1 receptor	[74]
	丝状真菌 Filamentous fungi	Pex7	PTS2受体 PTS2 receptor	[71,73]
	酵母 Yeast	Pex7	PTS2受体 PTS2 receptor	[74]
	丝状真菌 Filamentous fungi	Pex8	货物放行,进口装配 Cargo release, import assembly	[71,73]
	酵母 Yeast	Pex8	货物放行,进口装配 Cargo release, import assembly	[74]
	丝状真菌 Filamentous fungi	Pex13,Pex14, Pex17,Pex33	受体启动位点的组成部分 Components of the receptor initiation site	[71,73]
	酵母 Yeast	Pex13,Pex14,Pex17, Pex14-Pex17,Pex33	受体启动位点的组成部分 Components of the receptor initiation site	[74]
	丝状真菌 Filamentous fungi	Pex15,Pex26	Pex1和Pex6的锚定蛋白 Anchoring proteins of Pex1 and Pex6	[71,73]
	酵母 Yeast	Pex15,Pex26	Pex1和Pex6的锚定蛋白 Anchoring proteins of Pex1 and Pex6	[74]
	丝状真菌 Filamentous fungi	Pex18,Pex20,Pex21	Pex7的辅助受体 Co-receptor for Pex7	[71,73]
	酵母 Yeast	Pex18,Pex20,Pex21	Pex7的辅助受体 Co-receptor for Pex7	[74]
	丝状真菌 Filamentous fungi	Pex22	Pex4的锚定蛋白 Anchoring protein for Pex4	[71,73]
	酵母 Yeast	Pex22	Pex4的锚定蛋白 Anchoring protein for Pex4	[74]
调节过氧化物酶体的大小和丰度 Regulation of the size and abundance of peroxisomes	丝状真菌 Filamentous fungi	Pex11家族 Pex11 family	Dnm1的膜延长和GTPase活化蛋白 Membrane extension and GTPase activation proteins of Dnm1	[71,73]
	酵母 Yeast	Pex11	Dnm1的膜延长和GTPase活化蛋白 Membrane extension and GTPase activation proteins of Dnm1	[74]
	丝状真菌 Filamentous fungi	—	—	[71,73]
	酵母 Yeast	Pex25	膜的延伸和重塑 Membrane extension and remodeling	[74]
	丝状真菌 Filamentous fungi	—	—	[71,73]
	酵母 Yeast	Pex27	对裂变产生负面影响 Negative impact on fission	[74]
	丝状真菌 Filamentous fungi	—	—	[71,73]
	酵母 Yeast	Pex23,Pex24, Pex28,Pex29, Pex30,Pex31,Pex32	形成包含网状同源结构域的复合体 Formation of complexes containing reticulated homology domains	[74]

类别 Form	物种 Specie	蛋白 Protein	功能 Function	参考文献 Reference
过氧化物酶体膜的生物起源 Biological origin of	丝状真菌 Filamentous fungi	Pex3	Pex19膜锚定器 Pex19 membrane anchor	[71,73]
peroxisomal membranes	酵母 Yeast	Pex3	Pex19膜锚定器 Pex19 membrane anchor	[74]
	丝状真菌 Filamentous fungi	Pex19	mPTS受体 mPTS receptor	[71,73]
	酵母 Yeast	Pex19	mPTS受体 mPTS receptor	[74]
基质蛋白导入 Matrix protein introduction	丝状真菌 Filamentous fungi	Pex1,Pex6	参与受体循环的AAA型ATP酶 AAA-type ATPases involved in the receptor cycle	[71,73]
	酵母 Yeast	Pex1,Pex6	参与受体循环的AAA型ATP酶 AAA-type ATPases involved in the receptor cycle	[74]
	丝状真菌 Filamentous fungi	Pex2,Pex10,Pex12	参与受体泛素化(连接酶)的环复合体成分 Components of the ring complex involved in receptor ubiquitination (ligase)	[71,73]
	酵母 Yeast	Pex2,Pex10,Pex12	参与受体泛素化(连接酶)的环复合体成分 Components of the ring complex involved in receptor ubiquitination (ligase)	[74]
	丝状真菌 Filamentous fungi	Pex4	参与受体泛素化的泛素结合酶 Ubiquitin-binding enzymes involved in receptor ubiquitination	[71,73]
	酵母 Yeast	Pex4	参与受体泛素化的泛素结合酶 Ubiquitin-binding enzymes involved in receptor ubiquitination	[74]
	丝状真菌 Filamentous fungi	Pex5,Pex9	PTS1受体 PTS1 receptor	[71,73]
	酵母 Yeast	Pex5	PTS1受体 PTS1 receptor	[74]
	丝状真菌 Filamentous fungi	Pex7	PTS2受体 PTS2 receptor	[71,73]
	酵母 Yeast	Pex7	PTS2受体 PTS2 receptor	[74]
	丝状真菌 Filamentous fungi	Pex8	货物放行,进口装配 Cargo release, import assembly	[71,73]
	酵母 Yeast	Pex8	货物放行,进口装配 Cargo release, import assembly	[74]
	丝状真菌 Filamentous fungi	Pex13,Pex14, Pex17,Pex33	受体启动位点的组成部分 Components of the receptor initiation site	[71,73]
	酵母 Yeast	Pex13,Pex14,Pex17, Pex14-Pex17,Pex33	受体启动位点的组成部分 Components of the receptor initiation site	[74]
	丝状真菌 Filamentous fungi	Pex15,Pex26	Pex1和Pex6的锚定蛋白 Anchoring proteins of Pex1 and Pex6	[71,73]
	酵母 Yeast	Pex15,Pex26	Pex1和Pex6的锚定蛋白 Anchoring proteins of Pex1 and Pex6	[74]
	丝状真菌 Filamentous fungi	Pex18,Pex20,Pex21	Pex7的辅助受体 Co-receptor for Pex7	[71,73]
	酵母 Yeast	Pex18,Pex20,Pex21	Pex7的辅助受体 Co-receptor for Pex7	[74]
	丝状真菌 Filamentous fungi	Pex22	Pex4的锚定蛋白 Anchoring protein for Pex4	[71,73]
	酵母 Yeast	Pex22	Pex4的锚定蛋白 Anchoring protein for Pex4	[74]
调节过氧化物酶体的大小和丰度 Regulation of the size and abundance of peroxisomes	丝状真菌 Filamentous fungi	Pex11家族 Pex11 family	Dnm1的膜延长和GTPase活化蛋白 Membrane extension and GTPase activation proteins of Dnm1	[71,73]
	酵母 Yeast	Pex11	Dnm1的膜延长和GTPase活化蛋白 Membrane extension and GTPase activation proteins of Dnm1	[74]
	丝状真菌 Filamentous fungi	—	—	[71,73]
	酵母 Yeast	Pex25	膜的延伸和重塑 Membrane extension and remodeling	[74]
	丝状真菌 Filamentous fungi	—	—	[71,73]
	酵母 Yeast	Pex27	对裂变产生负面影响 Negative impact on fission	[74]
	丝状真菌 Filamentous fungi	—	—	[71,73]
	酵母 Yeast	Pex23,Pex24, Pex28,Pex29, Pex30,Pex31,Pex32	形成包含网状同源结构域的复合体 Formation of complexes containing reticulated homology domains	[74]

种类 Specie	PEX11基因数量 Number of PEX11 genes	PEX11基因种类 PEX11 gene type	功能 Function	参考文献 Reference
酵母类真菌 Yeast-like fungi	1	PEX11p	PEX11p缺失会导致过氧化物酶体数量减少,体积变大;过表达则会导致数量增多,体积减小 PEX11p deletion leads to a decrease in the number and size of peroxisomes; Overexpression leads to an increase in number and a decrease in size	[83]
人类、小鼠等哺乳动物 Humans, mice and other mammals	3	PEX11α	负责对外部刺激的过氧化物酶体增殖 Responsible for peroxisome proliferation in response to external stimuli	[1]
		PEX11β	组成型过氧化物酶体增殖所必需的 Necessary for the proliferation of the constitutive peroxisome	[1]
		PEX11γ	过氧化物酶体增殖所必需的,并与至少一种其他Pex11蛋白协同作用突出过氧化物酶体膜 Required for peroxisome proliferation and acts synergistically with at least one other Pex11 protein to highlight the peroxisome membrane	[87]
拟南芥、水稻 Arabidopsis thaliana, rice	5	PEX11ap	促进过氧化物酶体的复制 Promoting peroxisome replication	[84]
		PEX11bp	促进过氧化物酶体的聚集 Promoting peroxisome aggregation	[84]
		PEX11cp	促进过氧化物酶体无裂变的延伸 Promoting fissionless extension of peroxisome	[84]
		PEX11dp	促进过氧化物酶体无裂变的延伸 Promoting fissionless extension of peroxisome	[84]
		PEX11ep	促进过氧化物酶体的复制 Promoting peroxisome replication	[84]
丝状真菌 Filamentous fungi	1~5	PEX11A	缺失会导致过氧化物酶体明显数量减少,体积增大,同时损害了过氧化物酶体和伏鲁宁体的分离,侵染力下降,致病性降低 The deletion leads to a marked reduction in the number and increase in the size of peroxisomes, as well as impairing the segregation of peroxisomes and voroninosomes, with a decrease in invasive power and reduced pathogenicity	[81]
		PEX11B	在假禾谷镰孢菌中缺失会导致细胞壁完整性受损,活性氧代谢和脂肪酸β-氧化在内的细胞代谢过程明显受阻,脱氧雪腐镰刀菌烯醇(DON)产生能力下降 Deficiency in Fusarium pseudograminearum results in impaired cell wall integrity, marked impairment of cellular metabolic processes including reactive oxygen species metabolism and fatty acid β-oxidation, and decreased deoxynivalenol (DON) production capacity	[82]
		PEX11C	脂质代谢能力下降 Decreased lipid metabolism	[81]

Mechanism of peroxisome proliferation in different organisms

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物种 Specie	蛋白 Protein	功能 Function	参考文献 Reference
哺乳动物 Mammal	ACBD5、VAP	过氧化物酶体膜的重塑 Remodeling of peroxisomal membranes	[62]
植物 Plant	Pex11ap-ep	过氧化物酶体膜的重塑 Remodeling of peroxisomal membranes	[75]
真菌(酵母) Fungi (yeast)	Pex11p	过氧化物酶体膜的重塑 Remodeling of peroxisomal membranes	[94-95]
哺乳动物 Mammal	MIRO-运动蛋白复合物、Pex11βp、Pex14p MIRO-motor protein complex, Pex11βp, Pex14p	过氧化酶体膜的伸长 Elongation of peroxisomal membranes	[62]
植物 Plant	Pex11ap-ep	过氧化酶体膜的伸长 Elongation of peroxisomal membranes	[75]
真菌(酵母) Fungi (yeast)	Pex11p	过氧化酶体膜的伸长 Elongation of peroxisomal membranes	[94-95]
哺乳动物 Mammal	DRP1、Fis1、Mff、MIRO-运动蛋白复合物、Pex11βp DRP1, Fis1, Mff, MIRO-motor protein complex, Pex11βp	过氧化物酶体膜的收缩和分裂 Contraction and division of peroxisomal membranes	[62,96]
植物 Plant	DRP3A、DRP3B、DRP5B、FIS1A、FIS1B、PMD1	过氧化物酶体膜的收缩和分裂 Contraction and division of peroxisomal membranes	[82⇓-84]
真菌(酵母) Fungi (yeast)	Dnm1、Fis1、Mdv1、Pex11p、Vps1	过氧化物酶体膜的收缩和分裂 Contraction and division of peroxisomal membranes	[94-95]
哺乳动物 Mammal	—	过氧化物酶体的遗传 Inheritance of the peroxisome	—
植物 Plant	—	过氧化物酶体的遗传 Inheritance of the peroxisome	—
真菌(酵母) Fungi (yeast)	Inp1p、Inp2p、Myo2p	过氧化物酶体的遗传 Inheritance of the peroxisome	[92-93]

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