Guo-Qing Song

Guo-Qing Song

Contact Me

Department of Horticulture

(517) 353-0384


Quick links: Teaching    Publications    Research

Joined Department

September 01, 2007


15% Teaching, 30% Reseach, 50% Outreach and Contract Services, and 5% Administration

Associate Director, Plant Biotechnology Resource & Outreach Center

Research Experience

2022.7-present: Professor and Associate Director, Plant Biotechnology Resource & Outreach Center, Michigan State University, USA.

2014-2022: Associate professor and Associate Director, Plant Biotechnology Resource & Outreach Center, Michigan State University, USA.

2007-2014: Assistant professor and Associate Director, Plant Biotechnology Resource & Outreach Center, Michigan State University, USA.

2002-2007: Research Associate, Plant Transformation Center, Michigan State University, USA.

2000-2002: Research Associate, Biotechnology Group, Mitsui Chemical, INC. Japan.

1999.2-1999.8: Laboratoire de Cryobiologie Végétale, Université P. Et M. Curie, Paris, France


CSS/HRT 451 (3 credits): Biotechnology Applications for Plant Breeding and Genetics

Research Interest

Improvement of horticultural crops using genomic, genetic, and biotechnology approaches

Basic research 

Deciphering molecular basis of chilling-mediated flowering in woody plant

• Using K-domains of MADS box genes to improve crop/fruit yield
• VcFT-mediated flowering of blueberry plants
• VcDDF1-mediated abiotic tolerances in blueberry plants
• VcSOC1-memidated growth and development of blueberry plants
• The role of phytohormones in blueberry flowering 

Applied research

Engineering new generation of genetically modified (GM) crops

Yield enhancement: identify and functional analysis of flowering pathway genes of blueberry (e.g. VcSOC1, VcFT, VcCAL1, VcFUL, VcTFL1, VcAP1, and VcLFY) for the purpose of high yield for fruit crops. 
•  Abiotic tolerance/resistance:  using intragenic approach to improve woody fruit crops for abiotic (cold, heat, and drought) tolerance.
Transgrafting: using hairpin RNA derived siRNAs produced from transgenic rootstock to modify non-transgenic scion cultivars grafted on transgenic rootstock.

New Biotechnological Tool Development

Stable transformation systems  

Reliable transformation systems depend on efficient gene delivery, effective selection, and efficient plant regeneration. My previous transformation efforts using either Agrobacterium-or Biolistic-mediated transformation yielded successful transformation systems for 27 plant species, including: alfalfa, apple, Arabidopsis, atropa, barley, blueberry, canola, celery, common bean, grape, happy tree, petunia, rice, rutabaga, sour cherry, soybean, strawberry, sweet cherry, sweet cherry root stocks, sweetpotato, switchgrass, tobacco, tomato, wild tobaccos, wheat. 

Gene editing tools

Grape and blueberry

Micropropagation systems

Sweet cherry, apple, blueberry, cherry rootstock, asparagus, hop, blackberry, raspberry, chestnut, strawberry, hosta, and tree peony.

Selected Publications

*Corresponding author


Song GQ. 2024. Award-winning Fruit Publication in ASHS Journals across the Decades (1986–2023) HortTechnology 34 (3), 283-286.

Song GQ. Liu Z., Zhong GY. 2024. Regulatory frameworks involved in the floral induction, formation and developmental programming of woody horticultural plants: a case study on blueberries. Frontiers in plant science. 15, 1336892.

Arro J., Yang Y, Song GQ, Cousins P., Liu Z., Zhong GY. 2024. Transcriptome analysis unveils a potential novel role of VvAP1 in regulating the developmental fate of primordia into either inflorescences or tendrils in grapevine. Fruit Research 1-10.


Song GQ. Award-winning fruit publication in ASHS journals across the decades (1986-2023). HortTechnology. Accepted.

Gao D., Song GQ, Omar AA. 2023. Editorial: Genetic approaches for crop yield enhancement. Frontiers in plant science, 14.

Song GQ, Carter BB, Zhong GY. 2023. Multiple transcriptome comparisons reveal the essential roles of FLOWERING LOCUS T in floral initiation and SOC1 and SVP in floral activation in blueberry. Frontiers in Genetics 14.

Jacobs M, Thompson S, Platts AE, Body MJA, Kelsey A, Saad A, Abeli P, Teresi SJ, Schilmiller A, Beaudry R, Feldmann MJ, Knapp SJ, Song GQ, Miles T, Edger PP. 2023. Uncovering genetic and metabolite markers associated with resistance against anthracnose fruit rot in northern highbush blueberry. Hortic Res-England


Song GQ, Urban G, Ryner JR, Zhong G. 2022. Gene editing profiles in 94 CRISPR-Cas9 expressing T0 transgenic tobacco lines reveal high frequencies of chimeric editing of the target gene. Plants 11 (24), 3494

Yang Y, Ke J, Han X, Wuddineh WA, Song GQ, Zhong G. 2022, Removal of a 10-kb Gret1 transposon from VvMybA1 of Vitis vinifera cv. Chardonnay. Horticulture Research 9.

Kalendar R, Orbovic V, Egea-Cortines M, Song GQ. 2022. Recent advances in plant genetic engineering and innovative applications.Frontiers in plant science, 3978

Edger PP, Iorizzo M, NV  ... 2022. There and back again; historical perspective and future directions for Vaccinium breeding and research studies. Horticulture research 9

Han X-y, Yang Y, Han X, Ryner JT, Ahmed EAH, Qi Y, Zhong G, Song GQ . 2022. CRISPR Cas9- and Cas12a-mediated gusA editing in transgenic blueberry. Plant Cell, Tissue and Organ Culture.


Han X, Wang D, Song GQ*. 2021. Expression of a maize SOC1 gene enhances soybean yield potential through modulating plant growth and flowering. Sci Rep. 11: 12758.

Song GQ*, Han X, Ryner JT, Thompson A, Wang K. 2021. Utilizing MIKC-type MADS-box protein SOC1 for yield potential enhancement in maize. Plant Cell Rep.

Song GQ*, Han X. 2021. K-Domain Technology: Constitutive Expression of a Blueberry Keratin-Like Domain Mimics Expression of Multiple MADS-Box Genes in Enhancing Maize Grain Yield. Frontiers in Plant Science. 12: 844 (

Danial GH, Ibrahim DA, Song GQ (2021) Agrobacterium-mediated transformation of two tomato cultivars (lycopersicon esculentum mill.) cv. Sandra and rocky. Iraqi Journal of Agricultural Sciences 52(3): 745-755


Lin, T., Zhou, R., Bi, B., Song, L., Chai, M.*, Wang, Q.*, and Song, GQ*. 2020. Analysis of a radiation-induced dwarf mutant of a warm-season turf grass reveals potential mechanisms involved in the dwarfing mutant. Sci Rep 10 (1) 1-14, 18913 (2020).

Song, GQ*, Chen, Q., Callow, P., Mandujano, M., Han, Xy., Cuenca, B., Bonito, G., Medina-Mora, C., Fulbright, D.W., and Guyer, D.E. 2020. Efficient Micropropagation of Chestnut Hybrids (Castanea spp.) Using Modified Woody Plant Medium and Zeatin Riboside. Horticultural Plant Journal.

Zhao D, Zhong G-y, Song G-Q*. 2020. Transfer of endogenous small RNAs between branches of scions and rootstocks in grafted sweet cherry trees. PLoS ONE 15(7): e0236376.

Song, GQ*, Han, X., Wiersma, A.T., Zong, X.J., Awale, H.E., and Kelly, J.D. 2020. Induction of competent cells for Agrobacterium tumefaciens-mediated stable transformation of common bean (Phaseolus vulgaris L.). PLoS ONE 15(3): e0229909.

Song, GQ*, Walwort, A., and Zhao, D. 2020. Transgenic plants produced with a k-domain, and methods and expression cassettes related thereto. Patent: US10570409B2

Song GQ* 2020.Vaccinium spp. Blueberry and Cranberry, in Biotechnology of Fruit and Nut Crops, 2nd Edition edited by Richard E. Litz, Fernando Pliego Alfaro and Jose Ignacio Hormaza, CABI, Wallingford, Oxon OX10 8DE, UK. Page 191-205.


Song GQ*, Prieto, H., and Orbovic, V. (2019) Agrobacterium-Mediated Transformation of Tree Fruit Crops: Methods, Progress and Challenges. Frontiers in Plant Science doi: 10.3389/fpls.2019.00226

Lin, T., Walworth, A., Zong, X., Danial, G.H., Tomaszewski, E.M., Callow, P., Han, X., Zaharia, L.I., Edger, P.P., Zhong, G., Song GQ*(2019) VcRR2 regulates chilling-mediated flowering through expression of hormone genes in a transgenic blueberry mutant. Horticulture Research (2019)6:96 DOI:10.1038/s41438-019-0180-0

Song, GQ*, Walworth, A, Lin, T, Chen Q, Xiumei Han, L. Irina Zaharia,and Zhong G*. (2019). VcFT-induced mobile florigenic signals in transgenic and transgrafted blueberries. Horticulture Research. (2019)6(1):1-15 DOI: 10.1038/s41438-019-0188-5

Lin, T., Chen, Q.,  Wichenheiser, R.Z. and Song GQ* (2019) Constitutive expression of a blueberry FLOWERING LOCUS T gene hastens petunia plant flowering. Scientia Horticulturae 253 (27):376-381.

Zong, X.,  Denler, B.J. Danial, G.H., Chang, Y., and Song GQ* (2019) Adventitious Shoot Regeneration and Agrobacterium tumefaciens-mediated Transient Transformation of Almond × Peach Hybrid Rootstock ‘Hansen 536’. HortScience 54(5):936-940.

Zong, X., Zhang, Y., Walworth, A., Tomaszewski, E.M., Callow, P, G.-y Zhong  and Song GQ* (2019) Constitutive Expression of an Apple FLC3-like Gene Promotes Flowering in Transgenic Blueberry under Nonchilling Conditions. International Journal of Molecular Science. 20, 2775; doi:10.3390/ijms20112775

Arro, J., Yang, Y, Song GQ, and Zhong G*. (2019) RNA-Seq Reveals New DELLA Targets and Regulation in Transgenic GA-Insensitive Grapevines. BMC Plant Biology. 19 (1): DOI: 10.1186/s12870-019-1675-4

Song GQ*. (2019) Response of Woody Plants to Low Temperature. In Stress Physiology of Woody Plants (Dai W. ed.). Taylor & Francis Group. DOI:10.1201/9780429190476-8

Marivi Colle, Courtney P. Leisner, Ching Man Wai, Shujun Ou, Kevin A. Bird, Jie Wang, Jennifer H. Wisecaver, Alan E. Yocca, Elizabeth I. Alger, Haibao Tang, Zhiyong Xiong, Pete Callow, Gil Ben-Zvi, Avital Brodt, Kobi Baruch, Thomas Swale, Lily Shiue, Song GQ, Kevin L. Childs, Anthony Schilmiller, Nicholi Vorsa, C. Robin Buell, Robert VanBuren, Ning Jiang, and Patrick P. Edger*. 2019. Haplotype-phased genome and evolution of phytonutrient pathways of tetraploid blueberry. GigaScience, giz012,


Song GQ* and Walworth AE. (2018) An invaluable transgenic blueberry for studying chilling-induced flowering in woody plant. BMC Plant Biology,18: 265.

Song GQ* and Chen QX. (2018) Overexpression of the MADS-box gene K-domain increases the yield potential of blueberry. Plant Science. 276: 22–31

Song GQ* and Chen Q (2018) Comparative transcriptome analysis of nonchilled, chilled, and late-pink bud reveals flowering pathway genes involved in chilling-mediated flowering in blueberry. BMC Plant Biology 18:98 doi:10.1186/s12870-018-1311-8

Zong X, Chen Q, Mohamed AN, Kang Y, Lang G, and Song GQ* (2018) Adventitious shoot regeneration and Agrobacterium tumefaciens-mediated transformation of leaf explants of sweet cherry (Prunus avium L.). The Journal of Horticultural Science and Biotechnology. doi:10.1080/14620316.2018.1470908

Walworth AE and Song GQ* (2018) The cold-regulated genes of blueberry and their response to overexpression of VcDDF1 in several tissues. International Journal of Molecular Science. 19,1553; doi:10.3390/ijms19061553

Chen Q., Song GQ*. (2018) Protocol for Agrobacterium-Mediated Transformation and Transgenic Plant Production of Switchgrass. In: Kumar S., Barone P., Smith M. (eds) Transgenic Plants. Methods in Molecular Biology, vol 1864. Humana Press, New York, NY. Springer (


Guo-qing Song* and Xuan Gao (2017) Transcriptomic changes reveal gene networks responding to the overexpression of a blueberry DWARF AND DELAYED FLOWERING 1 gene in transgenic blueberry plants. BMC Plant Biology 17:106 DOI: 10.1186/s12870-017-1053-z.


Xuan Gao, Aaron Walworth, Charity Mackie, and Guo-qing Song* (2016) Overexpression of a blueberry FLOWERING LOCUS T is associated with changes in the expression of phytohormone related genes in blueberry plants. Horticulture Research doi:  10.1038/hortres.2016.53.

Aaron Walworth, Benli Chai, Guo-qing Song* 2016. Transcript profile of flowering regulatory genes in VcFT-overexpressing blueberry plants. PLOS ONE 11(6): e0156993. doi:10.1371/journal.pone.0156993.

Duan W, Wang, Liangsheng, Guo-qing Song* 2016. Agrobacterium tumefaciens-mediated transformation of wild tobacco species Nicotiana debneyi, Nicotiana clevelandii, and Nicotiana glutinosa. American Journal of Plant Science. 7, 1-7. doi: 10.4236/ajps.2016.71001.


Guo-qing Song*, Aaron Walworth, Wayne H. Loescher (2015) Grafting of genetically engineered plants. J. Amer. Soc. Hort. Sci. 140(3)203-213.

Guo-qing Song* (2015) Blueberry (Vaccinium corymbosum L.). In Agrobacterium Protocols (3 rd Edition)-Methods in Molecular Biology 1224 (Wang K. et al. ed.), Humana, Totowa, NJ. page 121-132.

Guo-qing Song* (2015) Cherry. In Agrobacterium Protocols (3 rd Edition)-Methods in Molecular Biology  1224 (Wang K. et al. ed.), Humana, Totowa, NJ. page 133-142.


Zhao Dongyan and Guo-qing Song* (2014) Rootstock-to-scion transfer of transgene-derived small interfering RNAs and their effect on virus resistance in non-transgenic sweet cherry. Plant Biotechnology Journal DOI: 10.1111/pbi.12243.

Zhao Dongyan and Guo-qing Song* (2014) High-throughput sequencing as an effective approach in profiling small RNAs derived from a hairpin RNA expression vector in woody plants. Plant Science. 228: 39-47.

Aaron Walworth, Guo-qing Song, Ryan M. Warner (2014) Ectopic AtCBF3 expression improves freezing tolerance and promotes compact growth habit in petunia. Molecular Breeding.  DOI 10.1007/s11032-013-9989-7

Guo-qing Song* (2014). Recent advances and opportunities in cherry biotechnology. Acta Horticulturae 1020:89-97. 

Lang, G.A., G.-Q. Song, K.C. Sink, A.E. Walworth, M.A. Cook, and R.F. Allison. (2014) Transgenic gene silencing confers increased resistance to Prunus necrotic ringspot virus in cherry rootstocks. Acta Hort. 1058:547-552.


Guo-qing Song*, Aaron Walworth, Dongyan Zhao, Britton Hildebrandt, and Michael Leasia. (2013) Constitutive expression of the K-domain of a Vaccinium corymbosum SOC1-like  (VcSOC1-K) MADS-box gene is sufficient to promote flowering. Plant Cell Reports (DOI: 10.1007/s00299-013-1495-1)

Guo-qing Song*, Aaron Walworth, Dongyan Zhao, Ning Jiang, and James F. Hancock. (2013)  The Vaccinium corymbosum FLOWERING LOCUS T-like gene (VcFT): a flowering activator reverses the photoperiodic and chilling requirements and enables early and continuous flowering in blueberry. Plant Cell Reports DOI: 10.1007/s00299-013-1489-z

Guo-qing Song*, Sink KC, Walworth AE, Cook MA, Allison RF, and Lang GA. 2013. Engineering cherry rootstocks with resistance to Prunus necrotic ring spot virus through RNAi–mediated silencing. Plant Biotechnology Journal  11(6):702-8 doi: 10.1111/pbi.12060      

Guo-qing Song* and Walworth AE. 2013. Agrobacterium tumefaciens-mediated transformation of Atropa belladonna. Plant Cell, Tissue and Organ Culture. Volume 115, Issue 1, pp 107-113,  DOI 10.1007/s11240-013-0342-y

Mukeshimana, G., Y. Ma, A. E. Walworth, Guo-qing Song, and J. D. Kelly. 2013. Factors influencing regeneration and Agrobacterium tumefaciens-mediated transformation of common bean (Phaseolus vulgaris L.). Plant Biotechnology Reports 7: 59-70. 


Walworth AE, Rowland LJ, Polashock JJ, Hancock JF and Guo-qing Song*. 2012. Overexpression of a blueberry-derived CBF enhances cold tolerance in a southern highbush blueberry cultivar. Molecular Breeding 30(3):1313-1323. DOI 10.1007/s11032-012-9718-7.

Guo-qing Song*, Walworth AE, and Hancock JF. 2012, Agrobacterium tumefaciens-mediated transformation of rutabaga (Brassica napus var. napobrassica) cultivar American Purple Top Yellow. In Vitro Cellular & Developmental Biology – Plant 48:383-389.

Guo-qing Song*, Walworth AE, and Hancock JF. 2012, Factors influencing Agrobacterium-mediated transformation of switchgrass cultivars. Plant Cell, Tissue and Organ Culture. 108(3): 445-453.

Guo-qing Song* and Hancock JF. 2012, Recent advances in blueberry transformation. International Journal of Fruit Science 12: 316-332.

Guo-qing Song*, Walworth AE, and Hancock JF. 2012, Stability of transgenes in blueberry. International Journal of Fruit Science 12: 333-341.


Guo-qing Song, Hancock J.F. 2011, Vaccinium. In Kole C (Ed) Wealth of Wild Crop Relatives: Genetic, Genomic & Breeding Resource. Springer-Verlag Berlin Heidelberg. P197-222.


Song GQ, Sink K.C., Hancock J.F. 2010, Flux in morphology and GUS expression in somatic embryogenic-derived populations of transgenic celery. In Kumar A (Ed) Applications of plant biotechnology: In vitro propagation, transformations, and secondary metabolite production. p 325-335.

Liu C, Callow P, Rowland LJ, Hancock JF, Song GQ*. 2010, Adventitious shoot regeneration from leaf explants of southern highbush blueberry cultivars. Plant Cell, Tissue and Organ Culture. 103: 137-144.

Song GQ, Sink KC, Ma Y, Herlache T, Hancock JF, Loescher WH. 2010, A novel mannose-based selection system for plant transformation using celery M6PR. Plant Cell Reports 29: 163-172


Song GQ, Sink K.C., Callow P.W., Baughan R., and Hancock, J.F. 2008, Evaluation of a herbicide resistant trait conferred by the bar gene driven by four distinct promoters in transgenic blueberry plants. Journal of the American Society for Horticultural Science 133(4): 605-611.

Loskutov AV, Song GQ, Sink KC. 2008, Evaluation of methods for celery (Apium Graveolens L.) transformation using Agrobacterium tumefaciens and the bar gene as selectable marker. In Vitro Cellular & Developmental Biology--Plant 44(4): 239-245.

Song GQ, Lang G., Dolgov S.V., and Sink K.C. 2008, Cherries. In: Kole C. and Hall T.C. (eds) A compendium of transgenic crop plants. Volume 4. p161-188. Wiley- Blackwell.

Hancock J.F., Song GQ, and Sink K.C. 2008, Berry Crops. In: Kole C. and Hall T.C. (eds) A compendium of transgenic crop plants. Volume 4. p93-120 Wiley- Blackwell. 


Song GQ and Yamaguchi K. 2007, Expression of a rice CAB promoter in sweetpotato plants. Journal of the American Society for Horticultural Science 132 (4): 551-556.

Song GQ, Roggers R.A., Sink K.C., Particka M., and Zandstra B. 2007, Production of herbicide-resistant highbush blueberry ‘Legacy’ by Agrobacterium-mediated transformation of the Bar gene. Acta Horticulturae 738: 397-408.

Song GQ and Sink K.C. 2007, Transformation of cherry: Prunus cerasus L. ‘Montmorency’ and Prunus cerasus x P. canescen ‘Gisela 6’ mediated by Agrobacterium tumefaciens and a two-step selection system. Acta Horticulturae 738: 683-690.

Song GQ and Sink K.C. 2007, Highly efficient Agrobacterium tumefaciens-mediated transformation of celery (Apium graveolens L.) through somatic embryogenic regeneration. Plant Cell, Tissue and Organ Culture 88: 193-200.


Song GQ and Sink K.C. 2006, Transformation and plant regeneration of Montmorency sour cherry (Prunus cerasus L.) and Gisela 6 (Prunus cerasus L. x P. canescens) cherry rootstock mediated by Agrobacterium tumefaciens. Plant Cell Rep. 25 (2): 117-123.

Song GQ and Sink K.C. 2006, Agrobacterium-mediated transformation of highbush blueberry (Vaccinium corymbosum L.) cultivars. In Agrobacterium Protocols (2th Edition)-Methods in Molecular Biology 344 (Wang K. et al. ed.), Humana, Totowa, NJ, pp 263-272.

Song GQ and Yamaguchi K. 2006, Agrobacterium tumefaciens-mediated transformation of sweet potato (Ipomoea batatas (L.) Lam.) using stem explants. In Agrobacterium Protocols (2th Edition)-Methods in Molecular Biology 344 (Wang K. et al. ed.), Humana, Totowa, NJ, pp 37-44.


Song GQ and Sink K.C. 2005, Optimizing adventitious shoot regeneration and Agrobacterium tumefaciens-mediated gene transfer system for sour cherry (Prunus cerasus L.) cultivar Montmorency. Scientia Horticulturae 106:60-69.

Song GQ and Sink K.C. 2005, Plant regeneration and genetic transformation of Blueberry. Recent Research Developments in Genetics & Breeding. Signpost press. Vol 2: 190-200.


Song GQ and Sink K.C. 2004, Agrobacterium tumefaciens-mediated transformation of blueberry (Vaccinium corymbosum L.) Plant Cell Rep. 23:475-484.

Song GQ, Honda H. and Yamaguchi K. 2004, Highly efficient Agrobacterium tumefaciens-mediated transformation of sweet Potato (Ipomoea batatas (L.) Lam.) from stem explants using a two-step kanamycin-hygromycin selection method. In Vitro Cellular & Developmental Biology--Plant 40:359-365.

Non-MSU related publications

Song GQ and Yamaguchi K, 2003, Efficient Agroinfiltration–mediated transient GUS expression system for assaying different promoters in rice. Plant Biotechnology, 20(3): 235-239.

Song GQ, Yu KS, Li SH, Liu GJ, Meng ZQ, and Zhu J, 2002, Forecasting fruit diameter of ‘Fuji’ apples (Malus Pumila Mill) at harvest during early growing season. Agricultural Sicences in China 1: 905-910.

Li SH, Song GQ, Liu GJ, Meng ZQ, Yu KS, and Zhu J, 2002, Forecasting of fruit size distribution of “Fuji” apples during the early growing season. Journal of Fruit Science 19:357-360.

Liu GJ, Song GQ, Li SH, and Meng ZQ, 2002, Effect of preharvest leaf-removal on apple fruit quality and branches reserve nutrient. China Fruits 2:11-13.

Bachiri Y, Song GQ, Plessis P, Shoar-Ghaffari A, Rekab T, and Morisset C, 2001, Routine crypreservation of kiwifruit (Actinidia spp) germplasm by encapsulation-dehydration: Importance of plant growth regulators. Cryo-letters 22:61-74.

Song GQ, Xiao XG, Li Sh H, Chen Sh W, and Gan M H. 2000, Expression of chicken Marek's Disease Virus B antigen in transgenic tobacco. Journal of Agricultural Biotechnology, 2:137-142.

Song GQ, Xiao XG, Li Sh H, 2000, Strawberry plant regeneration from leaf disks and Petioles, Deciduous Fruit 2: 3-5.

Related Work

All Related Work