闽楠紫色酸性磷酸酶基因（PAPs）家族分析及3个PAPs在低磷胁迫下的功能

doi:10.11707/j.1001-7488.LYKX20240323

摘要/Abstract

摘要：

目的: 紫色酸性磷酸酶（PAPs）是应对低磷胁迫的重要酸性磷酸酶，筛选响应低磷胁迫的闽楠PAPs基因，并进行功能验证，探究闽楠适应低磷胁迫下的响应机制，为培育闽楠磷高效利用种质提供理论依据。方法: 基于闽楠全基因组数据，通过blastp同源比对及hmmersearch对闽楠PAPs基因家族进行鉴定及生信分析，与拟南芥和挪威云杉PAPs蛋白序列比对，构建系统进化树。通过对闽楠2个家系（靖安3号和宜丰5号）苗在不同磷含量（0 mmol·L^?1 KH₂PO₄和1 mmol·L^?1 KH₂PO₄）处理下的光合荧光参数、无机磷含量、酸性磷酸酶（ACP）活性及PAPs表达量进行分析，筛选了3个PbPAPs进行后续基因功能研究。结果: 在闽楠全基因组中鉴定到22个PbPAPs基因，均含有Metallophos结构域，不均匀分布于10条染色体，氨基酸长度326~648 aa，大部分基因等电点小于7，外显子数量2~13个，共有2对串联重复和4对片段重复，系统进化树分为3组。低磷胁迫下2个家系苗主根伸长抑制，根毛密度和根表面积增加；净光合速率和F_v/F_m值均显著下降，而ACP活性显著升高；靖安3号叶片无机磷含量先下降后有所上升，宜丰5号变化较平稳，靖安3号处理7天后根无机磷含量显著下降，宜丰5号处理21天根无机磷含量显著下降，而处理后期变化趋势相近。经RT-qPCR分析，低磷诱导PbPAP10a、PbPAP15a和PbPAP26基因表达。分别通过发根农杆菌和根癌农杆菌介导遗传转化获得PbPAP10a、PbPAP15a和PbPAP26基因的过表达闽楠根系及转基因拟南芥，其ACP活性显著增强，无机磷含量显著升高，表明过表达PbPAPs基因可提高ACP活性和无机磷含量。结论: 闽楠对低磷胁迫有较高耐受性，且宜丰5号比靖安3号对低磷胁迫的耐受性更高，适用于瘠薄林地或酸性低磷林地造林，过表达闽楠PAPs基因可提高ACP活性和无机磷含量，助力于闽楠对低磷环境的适应性。

关键词: 闽楠, 紫色酸性磷酸酶, 低磷胁迫, 基因家族, 无机磷含量

Abstract:

Objective: Phoebe bournei is well adapted in red soils with low available phosphorus. Purple acid phosphatases (PAPs) are important acid phosphatases that hydrolyse a wide range of substrates in response to low phosphorus stress to release soluble inorganic phosphorus. This study aims to identify PAPs genes in response to low phosphorus stress and verify the functions in order to explore the response mechanism of P. bournei under low phosphorus stress, which will provide a theoretical basis for cultivating germplasm with efficient phosphorus utilization in P. bournei. Method: Based on the genome data of P. bournei, the PAPs gene family was identified by blastp homology and hmmersearch, and the protein sequences of PbPAPs were compared with those of Arabidopsis thaliana and Picea abies to construct a phylogenetic tree. The photosynthetic fluorescence parameters, inorganic phosphorus content, acid phosphatase (ACP) activity and the expression levels of PbPAPs under different phosphorus contents (0 mmol·L^?1 KH₂PO₄ and 1 mmol·L^?1 KH₂PO₄) in two P. bournei families (Jing'an No.3 and Yifeng No.5) were analyzed, and then three PbPAPs were screened out for the subsequent functional study. Result: A total of 22 PbPAPs genes were identified, all contained Metallophos domains, and were unevenly distributed on 10 chromosomes of P. bournei, with amino acid lengths ranging from 326 to 648 aa. Most of PbPAPs proteins had isoelectric points less than 7, and the exons number ranged from 2 to 13. These PbPAPs harbored two pairs of tandem duplications and four pairs of segmental duplications, and the phylogenetic tree was divided into 3 groups. Under low phosphorus treatments, the elongation of the primary root was inhibited, while the root hair density and the surface area of the root system were increased in the seedlings of P. bournei two families. The net photosynthetic rate and F_v/F_m value were decreased significantly, while the acid phosphatase activity increased. The inorganic phosphorus content of Jing'an No.3 leaves firstly declined and then increased slightly, while the change in Yifeng No.5 was relatively stable. A significant decrease of Pi content was observed in the Jing'an No.3 root after 7 days of treatment, while a significant decrease was firstly observed in the Yifeng No.5 after 21 days of treatment, and the change trends in the later stage of treatments were similar. According to RT-qPCR analysis, low phosphorus induced the gene expression level of PbPAP10a, PbPAP15a and PbPAP26. The three genes of PbPAP10a, PbPAP15a and PbPAP26 were overexpressed in P. bournei roots and A. thaliana by Agrobacterium tumefaciens- and A. rhizocarcinosum-mediated genetic transformation, which resulted in a significant enhancement of ACP activity and a significant increase in Pi content. Thus, overexpression of the PbPAPs gene increased ACP activity and Pi content. Conclusion: It is shown that P. bournei has high tolerance to low phosphorus stress, and Yifeng No.5 is more tolerant to low phosphorus stress than that of Jing'an No.3, which is suitable for forestation in barren forest land or acidic low phosphorus forest land. Overexpression of PbPAPs gene increases ACP activity and Pi content, which would help P. bournei to adapt to the low phosphorus environments.

Key words: Phoebe bournei, purple acid phosphatase, low phosphorus stress, gene family, Pi content

中图分类号:

S722.8

张漪曼,路龙俊,张毓婷,杨琪,韩潇,童再康,张俊红. 闽楠紫色酸性磷酸酶基因（PAPs）家族分析及3个PAPs在低磷胁迫下的功能[J]. 林业科学, 2025, 61(5): 146-160.

Yiman Zhang,Longjun Lu,Yuting Zhang,Qi Yang,Xiao Han,Zaikang Tong,Junhong Zhang. Analysis of Purple Acid Phosphatases Genes (PAPs) Family in Phoebe bournei and the Functions of Three PAPs under Low Phosphorus Stress[J]. Scientia Silvae Sinicae, 2025, 61(5): 146-160.

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成分Component	加磷 +Pi/(mmol·L^?1)	缺磷 -Pi/(mmol·L^?1)
CaCl₂·2H₂O	0.5	0.5
MgSO₄·7H₂O	0.5	0.5
KCl	0	1.0
KH₂PO₄	1.0	0
H₃BO₃	0.1	0.1
MnCl₂	20×10^?3	20×10^?3
CuSO₄·5H₂O	1×10^?3	1×10^?3
ZnSO₄·7H₂O	3×10^?3	3×10^?3
Na₂MoO₄·2H₂O	1×10^?3	1×10^?3
CoCl₂·6H₂O	1×10^?4	1×10^?4
FeSO₄·7H₂O	0.1	0.1

引物名称Primer	序列Sequence（5’→3’）
PbPAP10a-F	CAGTTGTGATGGGAGTGATGA
PbPAP10a-R	GTGAACCTGTTGTGGAGCAT
PbPAP15a-F	AAAGACCATCCCTACCACCC
PbPAP15a-R	GAGAGGGTGACGGAGATCTG
PbPAP26-F	CTTTTGAGCTCTGTGGGCAA
PbPAP26-R	ACCTTGAGTAATATGGACCTGCT
PbCBP20-F	GAAGGCAGACAATGGG
PbCBP20-R	ATAGTGCGATGGGACAG

引物名称Primer	序列Sequence（5’→3’）
PbPAP10a-F	ATGGGTTCTCCAGTTGTGATG
PbPAP10a-R	TTATGAATACGATTTGGTGGATTCATT
PbPAP15a-F	ATGGATTGGGATTTTCTCTCATTTG
PbPAP15a-R	CTACGCTGGGAAATGCAATT
PbPAP26-F	ATGGTAGAGGCTTATAGCAT
PbPAP26-R	TCAGAATGTGGCAATGCG

蛋白 Protein	基因编号 Gene ID	氨基酸 Amino acid	分子量 MW/kDa	等电点 PI
PbPAP1	Phbou.01G1223.1	613	69.05	6.10
PbPAP2	Phbou.01G1352.1	646	72.98	5.46
PbPAP9	Phbou.01G1351.1	648	73.94	6.16
PbPAP10a	Phbou.01G2906.1	473	54.53	6.55
PbPAP10b	Phbou.12G1471.1	456	52.32	6.70
PbPAP15a	Phbou.01G3123.1	545	62.01	5.51
PbPAP15b	Phbou.08G1918.1	547	61.92	5.26
PbPAP16a	Phbou.01G3288.1	376	42.08	7.13
PbPAP16b	Phbou.01G3289.1	353	39.96	6.00
PbPAP17a	Phbou.01G3733.1	326	36.73	5.43
PbPAP17b	Phbou.02G2250.1	329	37.38	5.25
PbPAP18	Phbou.10G0372.1	437	49.62	5.59
PbPAP20	Phbou.05G2290.1	422	46.90	5.58
PbPAP22	Phbou.04G1339.1	441	49.96	5.89
PbPAP23	Phbou.11G1361.1	543	60.84	5.52
PbPAP26	Phbou.03G0433.1	480	55.30	6.47
PbPAP27a	Phbou.01G3074.1	615	69.27	5.85
PbPAP27b	Phbou.08G1858.1	620	69.73	6.06
PbPAP27c	Phbou.03G0971.1	626	69.75	5.28
PbPAP28	Phbou.03G0037.1	402	45.51	5.80
PbPAP29a	Phbou.08G0566.1	606	66.91	8.49
PbPAP29b	Phbou.07G1718.1	372	41.02	5.36

蛋白 Protein	保守结构域 Domain
PbPAP1	Pur_ac_phosph_N	Metallophos	Metallophos_C
PbPAP2	Pur_ac_phosph_N	Metallophos	Metallophos_C
PbPAP9	Pur_ac_phosph_N	Metallophos	Metallophos_C
PbPAP10a	Pur_ac_phosph_N	Metallophos	Metallophos_C
PbPAP10b	Pur_ac_phosph_N	Metallophos	Metallophos_C
PbPAP15a	Pur_ac_phosph_N	Metallophos	Metallophos_C
PbPAP15b	Pur_ac_phosph_N	Metallophos	Metallophos_C
PbPAP16a		Metallophos
PbPAP16b		Metallophos
PbPAP17a		Metallophos
PbPAP17b		Metallophos
PbPAP18	Pur_ac_phosph_N	Metallophos	Metallophos_C
PbPAP20	Pur_ac_phosph_N	Metallophos	Metallophos_C
PbPAP22	Pur_ac_phosph_N	Metallophos	Metallophos_C
PbPAP23	Pur_ac_phosph_N	Metallophos	Metallophos_C
PbPAP26	Pur_ac_phosph_N	Metallophos	Metallophos_C
PbPAP27a	Pur_ac_phosph_N	Metallophos	Metallophos_C
PbPAP27b	Pur_ac_phosph_N	Metallophos	Metallophos_C
PbPAP27c	Pur_ac_phosph_N	Metallophos	Metallophos_C
PbPAP28		Metallophos
PbPAP29a		Metallophos
PbPAP29b		Metallophos