[1]

Malhotra H, Vandana, Sharma S, Pandey R. 2018. Phosphorus nutrition: plant growth in response to deficiency and excess. In Plant Nutrients and Abiotic Stress Tolerance, eds. Hasanuzzaman M, Fujita M, Oku H, Nahar K, Hawrylak-Nowak B. Singapore: Springer Singapore. pp. 171−190 doi: 10.1007/978-981-10-9044-8_7
[2]

Singh B, Satyanarayana T. 2011. Microbial phytases in phosphorus acquisition and plant growth promotion. Physiology and Molecular Biology of Plants 17:93−103

doi: 10.1007/s12298-011-0062-x
[3]

Shen J, Yuan L, Zhang J, Li H, Bai Z, et al. 2011. Phosphorus dynamics: from soil to plant. Plant Physiology 156:997−1005

doi: 10.1104/pp.111.175232
[4]

Tran HT, Hurley BA, Plaxton WC. 2010. Feeding hungry plants: the role of purple acid phosphatases in phosphate nutrition. Plant Science 179:14−27

doi: 10.1016/j.plantsci.2010.04.005
[5]

Bhadouria J, Giri J. 2022. Purple acid phosphatases: roles in phosphate utilization and new emerging functions. Plant Cell Reports 41:33−51

doi: 10.1007/s00299-021-02773-7
[6]

Schenk G, Mitić N, Hanson GR, Comba P. 2013. Purple acid phosphatase: a journey into the function and mechanism of a colorful enzyme. Coordination Chemistry Reviews 257:473−482

doi: 10.1016/j.ccr.2012.03.020
[7]

Mehra P, Giri J. 2020. Purple acid phosphatases (PAPs): molecular regulation and diverse physiological roles in plants. In Protein Phosphatases and Stress Management in Plants: Functional Genomic Perspective, ed. Pandey GK. Cham: Springer International Publishing. pp. 29−51 doi: 10.1007/978-3-030-48733-1_3
[8]

Wang L, Liu D. 2018. Functions and regulation of phosphate starvation-induced secreted acid phosphatases in higher plants. Plant Science 271:108−116

doi: 10.1016/j.plantsci.2018.03.013
[9]

Li D, Zhu H, Liu K, Liu X, Leggewie G, et al. 2002. Purple acid phosphatases of Arabidopsis thaliana. Journal of Biological Chemistry 277:27772−27781

doi: 10.1074/jbc.M204183200
[10]

Zhang Q, Wang C, Tian J, Li K, Shou H. 2011. Identification of rice purple acid phosphatases related to posphate starvation signalling. Plant Biology 13:7−15

doi: 10.1111/j.1438-8677.2010.00346.x
[11]

Zhu S, Chen M, Liang C, Xue Y, Lin S, et al. 2020. Characterization of purple acid phosphatase family and functional analysis of GmPAP7a/7b involved in extracellular ATP utilization in soybean. Frontiers in Plant Science 11:661

doi: 10.3389/fpls.2020.00661
[12]

González-Muñoz E, Avendaño-Vázquez AO, Montes RAC, de Folter S, Andrés-Hernández L, et al. 2015. The maize (Zea mays ssp. mays var. B73) genome encodes 33 members of the purple acid phosphatase family. Frontiers in Plant Science 6:341

doi: 10.3389/fpls.2015.00341
[13]

Hou L, Zhang D, Wu Q, Gao X, Wang J. 2025. Analysis and profiling of the purple acid phosphatase gene family in wheat (Triticum aestivum L). Protoplasma 262:73−86

doi: 10.1007/s00709-024-01983-6
[14]

Pang X, Cheng Y, Ruan M, Ye Q, Wang R, et al. 2022. The PAP gene family in tomato: comprehensive comparative analysis, phylogenetic relationships and expression profiles. Plants 11:563

doi: 10.3390/plants11040563
[15]

Yin C, Wang F, Fan H, Fang Y, Li W. 2019. Identification of tea plant purple acid phosphatase genes and their expression responses to excess iron. International Journal of Molecular Sciences 20:1954

doi: 10.3390/ijms20081954
[16]

Liu HC, Rao L, Meng JH, Zuo WT, Sun TT. 2025. Genome-wide analysis, identification, and transcriptional profile of the response to abiotic stress of the purple acid phosphatases (PAP) gene family in apple. International Journal of Molecular Sciences 26:1011

doi: 10.3390/ijms26031011
[17]

Zhou M, Chen W, Zhao M, Li Y, Li M, et al. 2021. Genome-wide characterization and evolutionary analyses of purple acid phosphatase (PAP) gene family with their expression profiles in response to low phosphorus stresses in Moso bamboo (Phyllostachys edulis). Forests 12:326

doi: 10.3390/f12030326
[18]

Li C, Liu J, Liao Y, Luo H, Wang L, et al. 2024. Genome-wide identification of the PAP gene family in potatoes and its expression response to phosphorus-solubilizing bacteria, freezing stresses, and phytohormonal regulation. Agronomy 14:2735

doi: 10.3390/agronomy14112735
[19]

Abbasi Vineh MA, Sabet MS, Karimzadeh G. 2021. Effect of AtPAP17 and AtPAP26 genes overexpression on yield and yield components under salt stress in Arabidopsis thaliana plant. Environmental Stresses in Crop Sciences 14:449−460

doi: 10.22077/escs.2020.2788.1725
[20]

Tarafdar S, Chowdhary G. 2025. Abiotic stress-inducible purple acid phosphatase from Solanum lycopersicum is targeted to peroxisome via non-canonical peroxisome targeting signal type 1. Acta Physiologiae Plantarum 47:57

doi: 10.1007/s11738-025-03800-w
[21]

O'Gallagher B, Ghahremani M, Stigter K, Walker EJL, Pyc M, Liu AY, et al. 2022. Arabidopsis PAP17 is a dual-localized purple acid phosphatase up-regulated during phosphate deprivation, senescence, and oxidative stress. Journal of Experimental Botany 73:382−399

doi: 10.1093/jxb/erab409
[22]

Ali Abbasi-Vineh M, Sabet MS, Karimzadeh G. 2021. Identification and functional analysis of two purple acid phosphatases AtPAP17 and AtPAP26 involved in salt tolerance in Arabidopsis thaliana plant. Frontiers in Plant Science 11:618716

doi: 10.3389/fpls.2020.618716
[23]

Wang B, Xie G, Liu Z, He R, Han J, et al. 2019. Mutagenesis reveals that the OsPPa6 gene is required for enhancing the alkaline tolerance in rice. Frontiers in Plant Science 10:759

doi: 10.3389/fpls.2019.00759
[24]

Guo ZY, Yin YH, Liu Y, Xie YT, Pei YH, et al. 2024. Cloning and expression pattern analysis of ZmPAP26b gene in maize under drought stress. Acta Agriculturae Boreali-Sinica 39:10−19

doi: 10.7668/hbnxb.20194777
[25]

El-Ramady H, Abdalla N, Kovacs S, Domokos-Szabolcsy E, Bákonyi N, et al. 2020. Sustainable biorefinery and production of alfalfa (Medicago sativa L): a review. Egyptian Journal of Botany 60:621−639

doi: 10.21608/ejbo.2020.37749.1532
[26]

Undersander D. 2021. Economic importance, practical limitations to production, management, and breeding targets of alfalfa. In The Alfalfa Genome, eds. Yu LX, Kole C. Cham: Springer International Publishing. pp. 1−11 doi: 10.1007/978-3-030-74466-3_1
[27]

Shen C, Du H, Chen Z, Lu H, Zhu F, et al. 2020. The chromosome-level genome sequence of the autotetraploid alfalfa and resequencing of core germplasms provide genomic resources for alfalfa research. Molecular Plant 13:1250−1261

doi: 10.1016/j.molp.2020.07.003
[28]

Hu Y, Qian C, Zhao X, Li J, Wang L, et al. 2024. Effects of nitrogen and phosphorus limitation on the growth and phosphorus allocation of alfalfa (Medicago sativa L). Journal of Soil Science and Plant Nutrition 24:343−353

doi: 10.1007/s42729-023-01541-8
[29]

Xiao K, Harrison M, Wang ZY. 2006. Cloning and characterization of a novel purple acid phosphatase gene (MtPAP1) from Medicago truncatula barrel medic. Journal of Integrative Plant Biology 48:204−211

doi: 10.1111/j.1744-7909.2006.00204.x
[30]

Chen C, Chen H, Zhang Y, Thomas HR, Frank MH, et al. 2020. TBtools: an integrative toolkit developed for interactive analyses of big biological data. Molecular Plant 13:1194−1202

doi: 10.1016/j.molp.2020.06.009
[31]

Duvaud S, Gabella C, Lisacek F, Stockinger H, Ioannidis V, et al. 2021. Expasy, the Swiss Bioinformatics Resource Portal, as designed by its users. Nucleic Acids Research 49:W216−W227

doi: 10.1093/nar/gkab225
[32]

Chou KC, Shen HB. 2010. Plant-mPLoc: a top-down strategy to augment the power for predicting plant protein subcellular localization. PLoS One 5:e11335

doi: 10.1371/journal.pone.0011335
[33]

Tamura K, Stecher G, Kumar S. 2021. MEGA11 molecular evolutionary genetics analysis version 11. Molecular Biology and Evolution 38:3022−3027

doi: 10.1093/molbev/msab120
[34]

Letunic I, Bork P. 2024. Interactive Tree of Life (iTOL) v6: recent updates to the phylogenetic tree display and annotation tool. Nucleic Acids Research 52:W78−W82

doi: 10.1093/nar/gkae268
[35]

Lescot M, Déhais P, Thijs G, Marchal K, Moreau Y, et al. 2002. PlantCARE, a database of plant cis-acting regulatory elements and a portal to tools for in silico analysis of promoter sequences. Nucleic Acids Research 30:325−327

doi: 10.1093/nar/30.1.325
[36]

O'Rourke JA, Fu F, Bucciarelli B, Yang SS, Samac DA, et al. 2015. The Medicago sativa gene index 1.2: a web-accessible gene expression atlas for investigating expression differences between Medicago sativa subspecies. BMC Genomics 16:502

doi: 10.1186/s12864-015-1718-7
[37]

Dong X, Deng H, Ma W, Zhou Q, Liu Z. 2021. Genome-wide identification of the MADS-box transcription factor family in autotetraploid cultivated alfalfa (Medicago sativa L) and expression analysis under abiotic stress. Bmc Genomics 22:603

doi: 10.1186/s12864-021-07911-9
[38]

Liu H, Li XY, He F, Wang X, Li MN, et al. 2024. Genome-wide identification of the SAUR gene family in medicago sativa and analysis of its expression pattern under abiotic stresses. Acta Prataculturae Sinica 33:135−153 (in Chinese)

doi: 10.11686/cyxb2023235
[39]

Schmittgen TD, Livak KJ. 2008. Analyzing real-time PCR data by the comparative CT method. Nature Protocols 3:1101−1108

doi: 10.1038/nprot.2008.73
[40]

Guo Q, Zhu S, Lai T, Tian C, Hu M, et al. 2026. A phosphate-starvation enhanced purple acid phosphatase, GmPAP23 mediates intracellular phosphorus recycling and yield in soybean. Plant, Cell & Environment 49:3871−3888

doi: 10.1111/pce.15400
[41]

Zeng XC, Yang YN, Li RY, Jiang XQ, Jiang X, et al. 2024. Identification of Alfalfa (Medicago sativa) MsCEP genes and functional analysis of its regulation in root growth and development. Scientia Agricultura Sinica 57:4839−4853 (in Chinese)

doi: 10.3864/j.issn.0578-1752.2024.24.002
[42]

Xu G, Guo C, Shan H, Kong H. 2012. Divergence of duplicate genes in exon–intron structure. Proceedings of the National Academy of Sciences of the United States of America 109:1187−1192

doi: 10.1073/pnas.1109047109
[43]

Yang Y, Zhou Y, Chi Y, Fan B, Chen Z. 2017. Characterization of soybean WRKY gene family and identification of soybean WRKY genes that promote resistance to soybean cyst nematode. Scientific Reports 7:17804

doi: 10.1038/s41598-017-18235-8
[44]

Durell SR, Ben-Naim A. 2017. Hydrophobic-hydrophilic forces in protein folding. Biopolymers 107:e23020

doi: 10.1002/bip.23020
[45]

Barberis E, Marengo E, Manfredi M. 2021. Protein subcellular localization prediction. In Proteomics Data Analysis, ed. Cecconi D. New York, NY: Springer US. pp. 197−212 doi: 10.1007/978-1-0716-1641-3_12
[46]

Masonbrink RE, Severin AJ, Seetharam AS. 2017. Comparative genomics of soybean and other legumes. In The Soybean Genome, eds. Nguyen HT, Bhattacharyya MK. Cham: Springer International Publishing. pp. 83−93 doi: 10.1007/978-3-319-64198-0_6
[47]

Li C, Gui S, Yang T, Walk T, Wang X, et al. 2012. Identification of soybean purple acid phosphatase genes and their expression responses to phosphorus availability and symbiosis. Annals of Botany 109:275−285

doi: 10.1093/aob/mcr246
[48]

Müller LM. 2021. Mapping membranes: lipid imaging reveals phosphorus-dependent membrane dynamics in Medicago truncatula nodules. Plant Physiology 185:1473−1475

doi: 10.1093/plphys/kiab039
[49]

Du PX, Liu H, Hu DX, Chen XP, Hong YB, et al. 2022. Genome-wide identification and expression analysis of purple acid phosphatase (PAP) gene family in peanut. Chinese Journal of Oil Crop Sciences 44:522−531 (in Chinese)

doi: 10.19802/j.issn.1007-9084.2021093
[50]

Dong S, Pang W, Liu Z, Li H, Zhang K, et al. 2022. Transcriptome analysis of leaf senescence regulation under alkaline stress in Medicago truncatula. Frontiers in Plant Science 13:881456

doi: 10.3389/fpls.2022.881456
[51]

Liao H, Wong FL, Phang TH, Cheung MY, Li WYF, et al. 2003. GmPAP3, a novel purple acid phosphatase-like gene in soybean induced by NaCl stress but not phosphorus deficiency. Gene 318:103−111

doi: 10.1016/S0378-1119(03)00764-9
[52]

Deng L, Chen F, Jiang L, Lam HM, Xiao G. 2014. Ectopic expression of GmPAP3 enhances salt tolerance in rice by alleviating oxidative damage. Plant Breeding 133:348−355

doi: 10.1111/pbr.12171
[53]

Reddy CS, Kim KM, James D, Varakumar P, Reddy MK. 2017. PgPAP18, a heat-inducible novel purple acid phosphatase 18-like gene (PgPAP18-like) from Pennisetum glaucum, may play a crucial role in environmental stress adaptation. Acta Physiologiae Plantarum 39:54

doi: 10.1007/s11738-017-2348-2
[54]

Xu H, Zhang H, Fan Y, Wang R, Cui R, et al. 2022. The purple acid phosphatase GmPAP17 predominantly enhances phosphorus use efficiency in soybean. Plant Science 320:111283

doi: 10.1016/j.plantsci.2022.111283
[55]

Bhadouria J, Singh AP, Mehra P, Verma L, Srivastawa R, et al. 2017. Identification of purple acid phosphatases in chickpea and potential roles of CaPAP7 in seed phytate accumulation. Scientific Reports 7:11012

doi: 10.1038/s41598-017-11490-9
[56]

Wang L, Lu S, Zhang Y, Li Z, Du X, et al. 2014. Comparative genetic analysis of Arabidopsis purple acid phosphatases AtPAP10, AtPAP12, and AtPAP26 provides new insights into their roles in plant adaptation to phosphate deprivation. Journal of Integrative Plant Biology 56:299−314

doi: 10.1111/jipb.12184