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Yan Zhao--Selected Publications
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A full list of publications can be found here


Pierro, R., Bottner-Parker, K., Panattoni, A., Wei, W., Marcone, C., Rizzo, D., Materazzi, A., Quaglino, F., Zhao, Y. (2023) Multilocus sequence typing of phytoplasmas associated with Flavescence dorée disease in Tuscany vineyards identifies a highly homogeneous lineage in the subgroup 16SrV-C. Crop Protection 163: DOI:

Wei, W., Shao, J., Bottner-Parker, K.D., and Zhao, Y. (2023) Draft genome sequence resource of CBPPT1, a ‘Candidatus Phytoplasma trifolii’-related strain associated with potato purple top disease in the Columbia Basin, USA. Plant Dis DOI:

Inaba, J., Shao, J., Trivellone, V., Zhao, Y., Dietrich, C.H., Bottner-Park, K., Ivanauskas, A. Wei, W. (2023) Guilt by association: DNA-barcoding based identification of potential plant hosts of phytoplasmas from their insect carriers Phytopathology PHYTO09220323-R. DOI:

Inaba, J., Kazeem, S.A., Zhao, Y., Zwolinska, A., Ogunfunmilayo, A.O., Arogundade, O., Wei, W. (2023) Tomato and jute mallow are two new hosts of papaya bunchy top phytoplasma, a 16SrXII-O subgroup strain in Nigeria. Plant Dis DOI: 10.1094/PDIS-09-22-2192-PDN.


Wei, W., Davis, R.E., Mowery, J. and Zhao, Y. (2022) Growth inhibition of phytopathogenic spiroplasmas by membrane-interactive antimicrobial peptides Novispirin T7 and Caerin 1.1. Ann Appl Biol 108:109-117. DOI:

Wheatley, M.S., Wang, Q., Wei, W., Bottner-Parker, K., Zhao, Y. and Yang, Y. (2022) Cas12a-based diagnostics for potato purple top disease complex associated with infection by ‘Candidatus Phytoplasma trifolii’-related strains. Plant Dis 106(8):2039-2045. DOI:

Wei, W. and Zhao, Y. (2022) Phytoplasma taxonomy: nomenclature, classification and identification. Biology 11(08):1119 DOI:

Wei, W., Inaba, J., Zhao, Y., Mowery, J.D., and Hammond, R. (2022) Phytoplasma infection blocks starch breakdown and triggers chloroplast degradation, leading to premature leaf senescence, sucrose reallocation, and spatiotemporal redistribution of phytohormones. Int J Mol Sci 23: 1810. DOI:


Zhao, Y., Wei, W., Davis, R.E., Lee, I.-M., Bottner-Parker, K.D. (2021) The agent associated with blue dwarf disease in wheat represents a new phytoplasma taxon, ‘Candidatus Phytoplasma tritici’. Int J Syst Evol Microbiol 105: DOI:

Kazeem, S.A., Inaba, J., Zhao, Y., Zwolinska, A., Ogunfunmilayo, A.O., Arogundade, O., Wei, W. (2021) Molecular identification and characterization of ‘Candidatus Phytoplasma convolvuli’-related strains (representing a new 16SrXII-O subgroup) associated with papaya bunchy top disease in Nigeria. Crop Protection 148: DOI:

Wei, W., Trivellone, V., Dietrich, C.H., Zhao, Y., Bottner-Parker, K.D., Ivanauskas, A. (2021) Identification of phytoplasmas representing multiple new genetic lineages from phloem-feeding leafhoppers highlights the diversity of phytoplasmas and their potential vectors. Pathogens 10:352.

Tan, Y., Li, Q., Zhao, Y., Wei, H., Wang, J., Baker, C.J., Liu, Q. and Wei, W. (2021) Integration of metabolomics and existing omics data reveals new insights into phytoplasma-induced metabolic reprogramming in host plants. PLOS ONE 16(2): e0246203.


Gasparich, G.E., Bertaccini, A., and Zhao, Y. (2020) Candidatus Phytoplasma In Bergey's Manual of Systematics of Archaea and Bacteria. Published by John Wiley & Sons, Inc. in association with Bergey’s Manual Trust. DOI:

Zhao, Y. and Wei, W. (2020) Datasheet report for ‘Candidatus Phytoplasma trifolii’ (clover proliferation phytoplasma). in Invasive Species Compendium. Wallingford, UK: CAB International.

Passera, A., Zhao, Y., Murolo, S., Roberto Pierro, R., Arsov, E., Mori, N., Moussa, A., Silletti, M.R., Casati, P., Panattoni, A., Wei, W., Mitrev, S., Materazzi, A., Luvisi, A., Romanazzi, G., Bianco, P.A., Davis, R.E., Quaglino, F. (2020) Multi-locus genotyping reveals new molecular markers for differentiating distinct genetic lineages among 'Candidatus Phytoplasma solani' strains associated with grapevine bois noir disease. Pathogens 9: 970. DOI:

Cai, W., Shao, J., Zhao, Y., Davis, R.E., and Costanzo, S. (2020) Draft genome sequence of ‘Candidatus Phytoplasma pini’-related strain MDPP: a resource for comparative genomics of gymnosperm-infecting phytoplasmas. Plant Dis 104:1009-1010. DOI:


Wei, W., Davis, R.E., Bauchan, G., and Zhao, Y. (2019) New symptoms identified in phytoplasma-infected plants reveal extra stages of pathogen-induced meristem fate-derailment. Molecular Plant-Microbe Interactions 32:1314-1323. DOI:

Tan, Y., Wang, J., Davis, R.E., Wei, H., Zong, X., Wei, W., Shao, J. Zhao, Y., and Liu, Q. (2019) Transcriptome analysis reveals a complex array of differentially expressed genes accompanying a source-to-sink changes in phytoplasma-infected sweet cherry leaves. Ann Appl Biol 175:69-82 DOI:

Pérez-López, E., Wei, W., Davis, R.E., Wang J., and Zhao, Y. (2019) First report of sugarcane yellow leaf disease in Mexico and detection of ‘Candidatus Phytoplasma asteris’-related strains in affected plants. Plant Dis. 103: DOI:

Wei, W., Zhao, Y., and Davis, R.E. (2019) Phytoplasma inoculum titer and inoculation timing influence symptom development in newly infected plants. Phytopathogenic Mollicutes 9: 115-116.

Davis, R.E., Shao, J.Y., Zhao, Y., Wei, W., Silver, A., Stump, Z., Gasparich, G.E., and Donofrio, N. (2019) Complete Genome Sequence of Spiroplasma phoeniceum strain P40T, a plant pathogen isolated from diseased plants of Madagascar periwinkle [Catharanthus roseus (L.) G. Don]. Genome Announc 8:e01612-18.  DOI: 10.1128/MRA.01612-18.

Davis, R.E., Dally, E.L., Zhao, Y., Webb, C. and Appel, A. (2019) First report of North American grapevine yellows (NAGY) in Kansas:  Association of ‘Candidatus Phytoplasma pruni’- and ‘Ca. Phytoplasma asteris’-related strains with the disease. Plant Dis 103: DOI: 10.1094/PDIS-05-18-0869-PDN.

Valiunas, D., Jomantiene, R., Ivanauskas, A., Sneideris, D., Zizyte-Eidetiene, M., Shao, J., Zhao, Y., Costanzo, S., and Davis, R.E. (2019) Rapid detection and identification of ‘Candidatus Phytoplasma pini’-related strains in Lithuania based on genomic markers present in 16S rRNA and tuf genes. Forest Pathology 00:e12553. DOI: 10.1111/efp.12553.


Wang, J., Liu, Q., Wei, W., Davis, R.E., Tan, Y., Lee, I.-M., Zhu, D., Wei, H. and Zhao, Y. (2018) Multilocus genotyping identifies a highly homogeneous phytoplasma lineage associated with sweet cherry virescence disease in china and its carriage by an erythroneurine leafhopper. Crop Protection 106: 13-22. DOI:

Davis, R.E., Dally, E.L., Zhao, Y. and Wolf, T.K. (2018) Genotyping points to divergent evolution of ‘Candidatus Phytoplasma asteris’ strains causing North American Grapevine Yellows and strains causing Aster yellows. Plant Dis 102: 1696-1702.

Davis, R.E. and Zhao, Y. (2018) Datasheet report for ‘Candidatus Phytoplasma pini’ (pine witches’-broom phytoplasma). in Invasive Species Compendium. Wallingford, UK: CAB International. DOI:


López, E.P., Wei, W., Wang, J., Davis, R.E., Rodríguez, M.L., and Zhao, Y. (2017) Novel phytoplasma strains of X-disease group unveil genetic markers that distinguish North American and South American geographic lineages within subgroups 16SrIII-J and 16SrIII-U. Ann Appl Biol 171: 405-416. DOI:

Naderali, N., Nejat, N., Vadamalai, G., Davis, R.E., Wei, W., Harrison, N.A., Kong, L., Kadir, J., Tan, Y.H. and Zhao, Y.  (2017) 'Candidatus Phytoplasma wodyetiae', a new taxon associated with yellow decline disease of foxtail palm (Wodyetia bifurcata). Int J Syst Evol Microbiol 67: 3765-3772. DOI:

Davis, R.E., Zhao, Y., Wei, W., Dally, E., and Lee, I.M. (2017) 'Candidatus Phytoplasma luffae', a novel taxon associated with a witches'-broom disease of loofah, Luffa aegyptica Mill. Int J Syst Evol Microbiol 67: 3127-3133. DOI:

Davis, R.E., Shao, J.Y., Zhao, Y., Gasparich, G.E., Gaynor, B.J. and Donofrio, N. Complete Genome Sequence of Spiroplasma citri strain R8-A2T, Causal Agent of Stubborn Disease in Citrus spp. (2017) Genome Announc 5(16): e00206-17.

Wei, W., Pérez-López, E., Davis, R.E., Bermúdez-Díaz, L., Granda-Wong, C., Wang, J., Zhao, Y. (2017) ‘Candidatus Phytoplasma brasiliense’-related strains associated with papaya bunchy top disease in northern Peru represent a distinct geographic lineage. Crop Protection 92: 99-106. DOI:

Quaglino, F., Murolo, S., Zhao, Y., Casati, P., Durante, G., Wei, W., Bianco, P.A., Romanazzi, G. Davis, R.E. (2017) Identification of new -J and -K 16SrXII subgroups and distinct single nucleotide polymorphism genetic lineages among ‘Candidatus Phytoplasma solani’ strains associated with bois noir in central Italy. Australian Plant Pathology 46: 31-34.


Wei, W., Pérez-López, E., Bermúdez-Díaz, L., Davis, R.E., Granda-Wong, C., Zhao, Y.  (2016) First report of a new grapevine yellows disease in Peru and its association with infection by a ‘Candidatus Phytoplasma brasiliense’-related phytoplasma strain. Plant Dis DOI:

Cai, H., Wang, L.C., Yang, Z.X., Wan, Q.L., Wei, W., Davis, R.E., Zhao, Y.  (2016) Evidence for the role of an invasive weed in widespread occurrence of phytoplasma diseases in diverse vegetable crops: implications from lineage-specific molecular markers. Crop Protection 89: 193-201. DOI:

Zhao, Y., Davis, R.E. (2016) Criteria for phytoplasma 16Sr group/subgroup delineation and the need of a platform for proper registration of new groups and subgroups. Int J Syst Evol Microbiol 66: 2121–2123. DOI:

Cai, H., Wang, L., Mu, W. Wan, Q., Wei, W., Davis, R.E., Chen, H., Zhao, Y. (2016) Multilocus genotyping of a ‘Candidatus Phytoplasma aurantifolia’-related strain associated with cauliflower phyllody disease in China. Ann Appl Biol 169: 64-74. DOI:

Costanzo, S., Rascoe, J., Zhao, Y., Davis, R.E., Nakhla, M. K. (2016). First report of a new ‘Candidatus Phytoplasma pini’-related strain associated with witches’-broom of Pinus spp. in Maryland. Plant Dis 100: 1776.

Wei, W., Wu, W., Davis, R.E., Lee, I-M., Zhao, Y. (2016) Development of molecular markers and a diagnostic tool for investigation of coinfections by and interactions between potato purple top and potato witches’-broom phytoplasmas in tomato. Ann Appl Biol 168: 133-141. DOI:

Davis, R.E., Harrison, N.A., Zhao, Y., Wei, W., and Dally, E.L. (2016) 'Candidatus Phytoplasma hispanicum', a novel taxon associated with Mexican periwinkle virescence disease of Catharanthus roseus. Int J Syst Evol Microbiol 66: 3463-3467. DOI:


Wei, W., Davis, R.E., Suo, X., Zhao, Y. (2015) Occurrence, distribution, and possible functional roles of simple sequence repeats in phytoplasma genomes. Int J Syst Evol Microbiol 65: 2748-2760. DOI:

Davis, R.E., Dally, E.L., Zhao, Y., Thomson, S.V., Nischwitz, C. (2015) Lilac (Syringa sp.) hosts a new ‘Ca. Phytoplasma pruni’-related strain in Utah: ribosomal RNA and secY genes distinguish the phytoplasma from previously described lineages. Plant Dis 100: 207.

Zhao, Y., Davis, R.E., Wei, W., Lee, I.-M. (2015) Should ‘Candidatus Phytoplasma’ be retained within the order AcholeplasmatalesInt J Syst Evol Microbiol 65: 1075-1082. DOI:

Davis, R.E., Dally L.E., Zhao, Y., Lee, I.-M., Wei, W., Wolf, T.K., Beanland, L., Ledoux, D.G., Johnson, D.A., and Fiola, J.A., Walter-Peterson, H., Dami, I., and Chien, M. (2015) Unraveling the etiology of north American grapevine yellows (NAGY): novel NAGY phytoplasma sequevars related to ‘Candidatus Phytoplasma pruni’. Plant Dis 99: 1087-1097. DOI:

Davis, R.E., Shao, J., Zhao, Y., Gasparich, G.E., Gaynor, B.J., and Donofrio, N. (2015) Complete genome sequence of Spiroplasma turonicum, a parasite of horse fly, Haematopota sp. (Diptera: Tabanidae). Genome Announc 3: e01367-15.

Davis, R.E., Shao, J., Dally, L.E., Zhao, Y., Gasparich, G.E., Gaynor, B.J., Athey, J.C., Harrison, N.A., and Donofrio, N. (2015) Complete genome sequence of Spiroplasma kunkelii strain CR2-3x, causal agent of corn stunt disease in Zea mays L.  Genome Announc 3: e01216-15.

Cheng, M., Dong, J., Lee, I.-M., Bottner-Parker, K., Zhao, Y., Davis, R.E., Laski, P.J., Zhang, Z., and McBeath, J.H. (2015) 16SrXII Group phytoplasma strains, including 16SrXII-E (‘Candidatus Phytoplasma fragariae’) and a new 16SrXII subgroup, 16SrXII-I, are associated with diseased potatoes (Solanum tuberosum) in the Yunan and Inner Mongolia regions of China. Eur J Plant Pathol 142: 305-318.

Lee, I.-M., Shao, J., Bottner-Parker, K., Gundersen D.E., Zhao, Y., and Davis, R.E. (2015) Draft genome sequence of ‘Candidatus Phytoplasma pruni’ strain CX, a plant pathogenic bacterium. Genome Announc 3: e01117-15.


Zhao, Y., Davis, R.E., Wei, W., Shao, J., and Jomantiene, R. (2014) Phytoplasma Genomes: evolution through mutually complementary mechanisms, gene loss and horizontal acquisition. In: Genomics of Plant-Associated Bacteria, Eds. Gross, D., Lichens-Park, A. & Kole, C., Springer-Verlag GmbH, Heidelberg pp234-271.

Chen, Y.P., Pettis, J.S., Corona, M., Chen, W.P., Li, C.J., Spivak, M., Visscher, P.K., DeGrandi-Hoffman, G., Boncristiani, H., Zhao, Y., van Engelsdorp, D., Delaplane, K., Solter, L., Drummond, F., Kramer, M., Lipkin, W.I., Palacios, G., Hamilton, M.C., Smith, B., Huang, S.K., Zheng, H.Q., Li, J.L. Zhang, X., Zhou, A.F., Wu L.Y., Zhou, J.Z., Lee, M.L., Teixeira, E.W., Li, Z.G., Evans, J.D. (2014) Israeli acute paralysis virus: epidemiology, pathogenesis and implications for honey bee health. PLoS Pathogens 10(7): e1004261. doi: 10.1371/journal.ppat.1004261.

Wang, J., Zhu, D., Liu, Q., Davis, R.E. and Zhao, Y.  (2014) First report of sweet cherry virescence disease in China and its association with infection by a ‘Candidatus Phytoplasma ziziphi’-related phytoplasma. Plant Dis 98: 419.

Lee, I.-M., Bottner-Parker, K., Zhao, Y., Davis, R.E., and Black, M.C. (2014) Aster yellows group (16Sr-I), subgroup 16SrI-A and 16SrI-B, phytoplasmas associated with lettuce yellows in Texas. New Dis Rep doi: 10.5197/j.2044-0588.2014.029.005.

Lee, I.-M., Polashock, J.J., Bottner-Parker, K., Bagadia, P.G., Rodriguez-Saona, C., Zhao, Y., and Davis, R.E. (2014) New subgroup 16SrIII-Y phytoplasmas associated with false-blossom diseases cranberry (Vaccinium macrocarpon) plants and with known and potential insect vectors in New Jersey. Eur J Plant Pathol 139: 399-406.

Li, J.L., Cornman, S.R., Evans, J.D., Pettis, J.S., Zhao, Y., Murphy, C., Peng, W.J., Wu, J., Hamilton, M., Boncristiani, H.F., Jr., Zhou, L., Hammond, J., and Chen,  Y. P. (2014) Invited Letter to the Editor (Reply to “Conclusive Evidence of Replication of a Plant Virus in Honeybees Is Lacking”) mBio 5(3) e01250-14. doi: 10.1128/mBio.01250-14.

Li, J., Cornman, S.R., Evans, J.D., Pettis, J.S., Zhao, Y., Murphy, C., Peng, W., Wu, J., Hamalton, M., Boncristiani, H.F., Zhou, L., Hammond, J. and Chen, Y. (2014) Systemic spread and propagation of a plant-pathogenic virus in European honeybees, Apis mellifera. mBio 5: e00898-13 doi: 10.1128/mBio.00898-13.


Wei, W., Davis, R.E., Nuss, D.L., and Zhao, Y. (2013) Phytoplasmal infection derails genetically preprogrammed meristem fate and alters plant architecture. Proc Natl Acad Sci USA 110: 19149-19154. DOI:

Chen, Y.P., Pettis, J.S., Zhao, Y., Liu, X., Tallon, L.J., Sadzewicz, L.D., Li, R., Zheng, H., Huang, S., Zhang, X., Hamilton, M.C., Pernal, S.F., Melathopoulos, A.P., Yan, X., and Evans, J.D. (2013) Genome sequencing and comparative genomics of honey bee microsporidia, Nosema apis reveal novel insights into host-parasite interactions. BMC Genomics 14: 451. doi: 10.1186/1471-2164-14-451.

Quaglino F, Zhao Y, Casati P, Bulgari D, Bianco PA, Wei, W., and Davis RE. (2013) ‘Candidatus Phytoplasma solani’, a novel taxon associated with stolbur and bois noir related diseases of plants. Int J Syst Evol Microbiol 63: 2879-2894. DOI:

Ding, Y., Wei, W., Wu, W., Davis, R.E., Jinag, Y., Lee, I.-M., Hammond, R.W., Shen, L., Sheng, J., and Zhao, Y. (2013) Role of gibberellic acid in tomato defense against potato purple top phytoplasma infection. Ann Appl Biol 162: 191–199. DOI:

Ding, Y., Wu, W., Wei, W., Davis, R.E., Lee, I.-M., Hammond, R.W., Sheng, J., Shen, L., Jinag, Y., and Zhao, Y. (2013) Potato purple top phytoplasma-induced disruption of gibberellin homeostasis in tomato plants. Ann Appl Biol 162: 131–139. DOI:

Bagadia, P.G., Polashock, J., Bottner-Parker K.D., Zhao, Y., Davis, R.E., and Lee, I.M. (2013) Characterization and molecular differentiation of 16SrI-E and 16SrIX-E phytoplasmas associated with blueberry stunt disease in New Jersey. Mol Cell Probes 27: 90-97.

Davis, R.E., Zhao, Y., Dally, E.L., Lee, I.-M., Jomantiene, R., and Douglas, S.M. (2013) ‘Candidatus Phytoplasma pruni’, a novel taxon associated with X-disease of stone fruits, Prunus spp.: multilocus characterization based on 16S rRNA, secY, and ribosomal protein genes. Int J Syst Evol Microbiol 63: 766-776. DOI:

Nejat, N., Vadamalai, G., Davis, R.E., Harrison, N.A., Sijam, K., Dickinson, M., Abdullah, S.N.A., and Zhao, Y. (2013) ‘Candidatus Phytoplasma malaysianum’, a novel taxon associated with virescence and phyllody of Madagascar periwinkle (Catharanthus roseus). Int J Syst Evol Microbiol 63: 540-548. DOI:


Zhao, Y., Wei, W., Lee, I.-M., Shao, J., Suo, X., and Davis, R.E. (2012) The iPhyClassifier, an interactive online tool for phytoplasma classification and taxonomic assignment. In Springer Protocols: Methods in Molecular Biology, Eds. Dickinson, M. & Hodgetts, J., Humana Press, New York, USA pp 329-338. DOI:

Lee, I.-M., Bottner-Parker, K.D., Zhao, Y., Bertaccini, A., and Davis, R.E. (2012) Differentiation and classification of phytoplasmas in the pigeon pea witches'-broom group (16SrIX): an update based on multiple gene sequence analysis. Int J Syst Evol Microbiol 62: 2279-2285. DOI:

Wu, W., Ding, Y., Wei, W., Davis, R.E., Lee, I.-M. Hammond, R.W., and Zhao, Y. (2012) Salicylic acid-mediated elicitation of tomato defense against infection by potato purple top phytoplasma. Ann Appl Biol 161: 36-45. DOI:

Wu, W., Cai, H., Wei, W., Davis, R.E., Lee, I.M., Chen, H., and Zhao, Y. (2012) Identification of two new phylogenetically distant phytoplasmas from Senna surattensis plants exhibiting stem fasciation and shoot proliferation symptoms. Ann Appl Biol 160:25-34. DOI:

Davis, R.E., Zhao, Y., Dally, E.L., Jomantiene, R., Lee, I.M., Wei, W., and Kitajima, E.W. (2012) 'Candidatus Phytoplasma sudamericanum', a novel taxon, and strain PassWB-Br4, a new subgroup 16SrIII-V phytoplasma, from diseased passion fruit (Passiflora edulis f. flavicarpa Deg.). Int J Syst Evol Microbiol 62: 984-989. DOI:


Wei, W., Cai, H., Jiang, Y., Lee, I.-M., Davis, R.E., Ding, Y., Yuan, E., Chen, H., and Zhao, Y. (2011) A new phytoplasma associated with little leaf disease in azalea: multilocus sequence characterization reveals a distinct lineage within the aster yellows phytoplasma group. Ann Appl Biol 158: 318-330. DOI:

Jiang, H., Wang, Y., Jiang Y., Wei, W., Jomantiene, R., Davis, R.E., and Zhao, Y. (2011) A PCR-based assay for rapid detection and precise differentiation of three quarantine phytoplasmas responsible for yellows diseases in grapevine.  Acta Phytopathologica Sinica 41: 352-360.

Jomantiene, R., Zhao, Y., Lee, I.-M., and Davis, R.E. (2011) Phytoplasmas infecting sour cherry and lilac represent two distinct lineages having closes evolutionary affinities with clover phyllody phytoplasma. Eur J Plant Pathol 130: 97-107.

Lee, I.-M., Bottner-Parker, K.D., Zhao, Y., Villalobos, W. and Moreira, L. (2011) ‘Candidatus Phytoplasma costaricanum’ a new phytoplasma associated with a newly emerging disease in soybean in Costa Rica. Int J Syst Evol Microbiol 61: 2822-2826.

Tuo, W., Zhao, Y., Zhu, D., and Jenkins, M.C. (2011) Immunization of female BALB/c mice with Neospora cyclophilin and/or NcSRS2 elicits specific antibody response and prevents against challenge infection by Neospora caninumVaccine 29: 2392-2399.

Kovalskaya, N., Zhao, Y., and Hammond, R.W. (2011) Antimicrobial and antifungal activity of a snakin-defensin hybrid protein expressed in tobacco and potato plants. Open Plant Sci J 5: 29-42.


Jomantiene, R., Davis, R.E., Lee, I.-M., Zhao, Y., Bottner-Parker, K., Valiunas, D., and Petkauskaite, R. (2010) Onion is a host for two phytoplasma lineages, subgroups 16SrI-A and 16SrI-(B/L)L, in Lithuania: a HinfI site revealed SNPs marking divergent branches of evolution. J Plant Pathol 92: 451-460.

Lee, I.-M., Bottner, K.D., Zhao, Y., Davis, R.E., and Harrison, N.  (2010) Phylogenetic analysis and delineation of phytoplasmas based on the secY gene.  Int J Syst Evol Microbiol 60: 2887-2897.

Zhao, Y., Wei, W., Davis, R.E. and Lee, I.-M. (2010) Recent advances in 16S rRNA gene-based phytoplasma differentiation, classification and taxonomy. In Phytoplasmas: Genomes, Plant Hosts and Vector, Eds. Weintraub, P. & and Jones, P., CABI Publishing, Wallingford, UK. pp 64-92.

Lee, I.-M., Zhao, Y. and Davis, R.E. (2010) Prospects of multiple gene-based systems for differentiation and classification of phytoplasmas. In Phytoplasmas: Genomes, Plant Hosts and Vector, Eds. Weintraub, P. & and Jones, P., CABI Publishing, Wallingford, UK. pp 51-63.

Davis, R.E., Dally, E.L., Zhao, Y., Lee, I.-M., Jomantiene, R., Detweiler, A.J., and Putnam, M.L.  (2010) First report of a new subgroup 16SrIX-E, ‘Candidatus Phytoplasma phoenicium’-related, phytoplasma associated with juniper witches’-broom disease in Oregon.  Plant Pathology 20: 35.


Zhao, Y., Sun, Q., Wei, W., Davis, R. E., Wu, W., and Liu, Q. (2009) ‘Candidatus Phytoplasma tamaricis’, a novel taxon discovered in witches’-broom-diseased salt cedar (Tamarix chinensis Lour.). Int J Syst Evol Microbiol 59: 2496-2504. DOI:

Zhao, Y., Wei, W., Lee, I.-M., Shao, J., Suo, X., and Davis, R. E. (2009) Construction of an interactive online phytoplasma classification tool, iPhyClassifier, and its application in analysis of the peach X-disease phytoplasma group (16SrIII). Int J Syst Evol Microbiol 59: 2582-2593. DOI:

Hammond, R.W. and Zhao, Y. (2009) Modification of tobacco plant development by sense and antisense expression of the tomato viroid-induced AGC VIIIa protein kinase PKV suggests involvement in gibberellin signaling. BMC Plant Biol 9: 108.

Quaglino, F., Zhao, Y., Bianco, P., Wei, W., Casati, P., Durante, G., and Davis, R.E. (2009) New 16Sr subgroups and distinct SNP lineages among grapevine Bois noir phytoplasma populations. Ann Appl Biol 154: 279-289.

Cornman, R.S., Chen, Y.P., Schatz, M.C., Street, C., Zhao, Y., Desany, B., Egholm, M., Hutchison, S., Pettis, J.S., Lipkin, W.I., and Evans, J.D. (2009) Genomic analyses of the microsporidian Nosema ceranae, an emergent pathogen of honey bees. PLoS Pathog 2009 5: e1000466.

Valiunas, D., Jomantiene, R., Ivanauskas, A., Abraitis, R., Staniene, G., Zhao, Y. and Davis, R.E. (2009) First report of a new phytoplasma subgroup, 16SrIII-T, associated with decline disease affecting sweet and sour cherry trees. Plant Dis 93: 500.


Wei, W., Davis, R.E., Jomantiene, R., and Zhao, Y.  (2008) Ancient, recurrent phage attacks and recombination events shaped dynamic sequence-variable mosaic structures at the root of phytoplasma genome evolution. Proc Natl Acad Sci USA 105: 11827-11832. DOI:

Wei, W., Lee, I-M., Davis, R.E., Suo, X., and Zhao, Y.  (2008) Automated RFLP pattern comparison and similarity coefficient calculation for rapid delineation of new and distinct phytoplasma 16S rDNA subgroup lineages. Int J Syst Evol Microbiol 58: 2368 – 2377. DOI:

Cai, H., Wei, W., Davis, R.E., Chen, H., and Zhao, Y. (2008) Genetic diversity among phytoplasmas infecting Opuntia species: virtual RFLP analysis identifies new subgroups in the peanut witches’-broom phytoplasma group. Int J Syst Evol Microbiol 58: 1448-1457. DOI:

Abraitiene, A., Zhao, Y., and Hammond, R. (2008) Nuclear targeting by fragmentation of the potato spindle tuber viroid genome. Biochem Biophys Res Commun 368: 470-475.


Wei, W., Davis, R.E., Lee, I.-M., and Zhao, Y. (2007) Computer-simulated RFLP analysis of 16S rRNA genes: identification of ten new phytoplasma groups. Int J Syst Evol Microbiol 57: 1855-1867. DOI:

Jomantiene, R., Davis, R.E., and Zhao, Y. (2007) Sequence variable mosaics: composites of recurrent transposition characterizing the genomes of phylogenetically diverse phytoplasmas.  DNA Cell Biol 26: 557-564.

Wei, W., Jiang, H., Yang, Y., Wang, Y., Davis, R.E., and Zhao, Y. (2007) Molecular identification of a new phytoplasma strain associated with the first observation of jujube witches’-broom disease in northeastern China. Plant Dis 91: 1364.

Lee, I.-M., Zhao, Y., Davis, R.E., Wei, W., and Martini, M. (2007) Prospects of DNA-based systems for differentiation and classification of phytoplasmas. Bull Insectol 60: 239-244.

Wei, W., Cai, H., Chen, H., Davis, R.E., and Zhao, Y. (2007) First report of a natural infection of Opuntia sp. by a ‘Candidatus Phytoplasma asteris’-related phytoplasma in China. Plant Dis 91: 461.

Martini, M., Lee, I.-M., Bottner, K. D., Zhao, Y., Botti, S., Bertaccini, A., Harrison, N. A., Carraro, L., Marcone, C., Khan, A. J., and Osler, R. (2007) Ribosomal protein gene-based phylogeny for finer differentiation and classification of phytoplasmas. Int J Syst Evol Microbiol 57: 2037-2051.

Zhao, Y., Sun, Q., Davis, R.E., Lee, I-M., and Liu, Q. (2007) First report of witches’-broom disease in a Cannabis species and its association with a phytoplasma of elm yellows group (16SrV). Plant Dis 91: 227.   


Wei, W., Opgenorth, D.C., Davis, R.E., Chang, C.J., Summers, C.G., and Zhao, Y. (2006) Characterization of a novel adhesin gene and design of a real-time PCR for rapid, sensitive and specific detection of Spiroplasma kunkeliiPlant Dis 90: 1233-1238. DOI:

Dally, E.L., Barros, T.S., Zhao, Y., Lin, S., Bruce, R.A., and Davis, R.E. (2006) Physical and genetic map of the Spiroplasma kunkelii CR2-3X chromosome.  Can J Microbiol 52: 857-867. 

Lee, I.-M., Zhao, Y., and Bottner, K.D. (2006) SecY gene sequence analysis for finer differentiation of diverse strains in the aster yellows phytoplasma groups.  Mol Cell Probes 20: 87-91.

Shao, J.Y., Jomantiene, R., Dally, E.L., Zhao, Y., Lee, I.-M., Nuss, D.L., and Davis, R.E. (2006) Phylogeny and characterization of phytoplasmal NusA and use of the nusA gene in detection of group 16SrI strains. J Plant Pathol 88: 189-197.


Zhao, Y., Davis, R.E., and Lee, I.-M. (2005) Phylogenetic positions of ‘Candidatus Phytoplasma asteris’ and Spiroplasma kunkelii as inferred from multiple sets of concatenated core house-keeping proteins. Int J Syst Evol Microbiol 55: 2131-2141. DOI:

Zhao, Y. and Hammond, R.W. (2005) Development of a candidate vaccine for Newcastle Disease Virus by epitope display in the Cucumber mosaic virus capsid protein.  Biotechnol Lett 27: 375-382.

Davis, R.E., Jomantiene, R., and Zhao, Y. (2005) Lineage-specific decay of folate biosynthesis genes suggests ongoing host adaptation in phytoplasmas. DNA Cell Biol 24: 832-840.

Liu, Q., Sun, Q., Wu, T., Davis, R.E., and Zhao, Y. (2005) First report of witches’-broom disease of Chinese wingnut (Pterocarya stenoptera) and its association with a phytoplasma of elm yellows group (16SrV).  Plant Dis 89: 529.

Lee, I.-M., Zhao, Y., and Bottner, K.D.  (2005) Novel insertion sequence-like elements in phytoplasma strains of the aster yellows group are putative new members of the IS3 family.  FEMS Microbiol Lett 242: 353-360.

Davis, R.E., Dally, E.L., Jomantiene, R., Zhao, Y., Roe, B.A., Lin, S., and Shao, J. (2005) Molecular analysis of a cryptic plasmid encoding a putative adhesin and type IV secretion system components in the wall-less plant pathogen Spiroplasma kunkeliiPlasmid 53: 179-190.


Zhao, Y., Liu, Q., and Davis, R.E. (2004) Transgene expression in strawberries driven by a heterologous phloem-specific promoter.  Plant Cell Rep 23: 224-230.

Zhao, Y., Hammond, R.W., Lee, I.-M., Roe, B.A., Lin, S., and Davis, R.E.  (2004) Cell division gene cluster in Spiroplasma kunkelii: functional characterization of ftsZ and the first report of ftsA in mollicutes.  DNA Cell Biol 23: 127-134.

Zhao, Y., Wang, H., Hammond, R.W., Jomantiene, R, Liu, Q., Lin, S., Roe, B.A., and Davis, R.E. (2004) Predicted ATP-binding cassette systems in the phytopathogenic mollicute Spiroplasma kunkelii.  Mol Genet Genomics 271: 325-338.

Liu, Q., Wu, T., Davis, R.E., and Zhao, Y.  (2004) First report of witches’-broom disease of Broussonetia papyrifera and its association with a phytoplasma of aster yellows group (16SrI).  Plant Dis 88: 770.

Chen, Y., Zhao, Y., Hammond, J., Hsu, H.T., Evans, J., and Feldlaufer, M. (2004) Multiple virus infection in the honey bee and genome divergence of honey viruses.  J Invertebr Pathol 87: 84-93.


Zhao, Y., Hammond, R.W., Jomantiene, R., Dally, E.L., Lee, I.-M., Jia, H., Wu, H., Lin, S., Zhang, P., Kenton, S., Najar, F.Z., Hua, A., Roe, B.A., Fletcher, J., and Davis, R.E. (2003) Gene content and organization of an 85-kb DNA segment from the genome of the phytopathogenic mollicute Spiroplasma kunkelii.  Mol Genet Genomics 269: 592-602.

Davis, R.E., Jomantiene, R., Zhao, Y., and Dally, E.L. (2003) Folate Biosynthesis pseudogenes, YfolP and YfolK, and an O-sialoglycoprotein endopeptidase gene homolog in the phytoplasma genome.  DNA Cell Biol 22: 697-706.


Zhao, Y., Owens, R.A., and Hammond, R.W. (2001) Use of a vector based on Potato virus X in a whole plant assay to demonstrate nuclear targeting of Potato spindle tuber viroid.  J Gen Virol 82: 1491-1497.


Zhao, Y., Hammond, J., Tousignant, M.E., and Hammond, R.W. (2000) Development and evaluation of a complementation-dependent gene delivery system based on Cucumber mosaic virus.  ArchVirol 145: 2285-2295.

Hammond, R.W. and Zhao, Y. (2000) Characterization of a tomato protein kinase gene induced by infection by Potato spindle tuber viroid.  Mol Plant-Microbe Interact 13: 903-910.