William Neal Reynolds Distinguished Professor

Center for Integrated Fungal Research
Department of Entomology and Plant Pathology,
223 Partners III, Box 7567
North Carolina State University
Raleigh, NC 27695-7567
Phone: 919.515.6994
Fax: 919.513.0024
Email :


B.S., North Carolina State University
M.S., Cornell University
Ph.D., Cornell University
Research Associate, Laboratory of Agricultural Biochemistry, Lincoln, NE


Visit my lab

Genomic approaches and field studies are being used in my program to improve food safety and to understand fungal biology and secondary metabolism. Gene expression profiling is being used to characterize the regulatory circuits in Aspergillus flavus governing aflatoxin biosynthesis and to elucidate the molecular response in maize kernels to Aspergilus flavus infection.. Field and laboratory studies are focused on understanding the ecology of Aspergillus flavus and Fusarium verticillioides and the inheritance of resistance to the fungi. Maize seed proteins inhibitory to mycotoxin-producing fungi are being isolated and characterized to better understand resistance mechanisms in seeds.


PP 707 Spring Semester
Fundamental concepts and current status of research on the physiology, biochemistry and molecular biology of host-pathogen interactions. Topics include recognition, penetration and colonization, pathogenicity and virulence determinants, resistance mechanisms, signal transduction, programmed cell death, and other current topics. Information presented in context of viral-, bacterial-, fungal-and nematode-plant interactions.

PP 730 Fall Semester
Basic concepts of genetics, ecology, and physiology of fungi, with emphasis on saprophytic and plant and animal pathogenic mycelial fungi. Current literature on evolution, cell structure, growth and development, gene expression, metabolism, sexual and asexual reproduction and incompatibility systems.

Selected Publications

Niu, C., Payne, G.A. and Charles P Woloshuk, C.P. 2015. Transcriptome changes in Fusarium verticillioides caused by mutation in the transporter-like gene FST1. BMC Microbiology. 15: 90. doi:10.1186/s12866-015-0427-3.

Shu, X., Livingston, D. P., III, Franks, R. G., Boston, R. S., Woloshuk, C. P. and Payne, G. A. 2014. Tissue-specific gene expression in maize seeds during colonization by Aspergillus flavus and Fusarium verticillioides. Mol Plant Pathol. doi: 10.1111/mpp.12224.

Dolezal, A. L., Shu, X. M., OBrian, G. R., Nielsen, D. M., Woloshuk, C. P., Boston, R. S., et al. 2014. Aspergillus flavus infection induces transcriptional and physical changes in developing maize kernels. Front Microbiol, 5: 384. doi:10.3389/fmicb. 2014.00384.

Reverberi, M., Punelli, M., Scala, V., Scarpari, M., Uva, P., Mentzen, W. I., Dolezal, A.L., Woloshuk, C., Pinzari, F., Fanelli, C. and Payne, G.A. 2013. Genotypic and phenotypic versatility of Aspergillus flavus during maize exploitation. Plos One, 8 (7).

Dolezal, A. L., Obrian, G. R., Nielsen, D. M., Woloshuk, C. P., Boston, R. S. and Payne, G. A. 2013. Localization, morphology and transcriptional profile of Aspergillus flavus during seed colonization. Mol Plant Pathol, 14: 898-909.

Reverberi, M., Punelli, M., Smith, C. A., Zjalic, S., Scarpari, M., Scala, V., Cardinali, G.,Aspite, N., Pinzari, F., Payne, G.A., Fabbri, A.A., Fanelli, C. 2012. How peroxisomes affect aflatoxin biosynthesis in Aspergillus flavus. Plos One, 7 (10).

Joardar, V., Abrams, N. F., Hostetler, J., Paukstelis, P. J., Pakala, S., Pakala, S. B., Zafar,N., Abolude, O.O., Payne, G., Andrianopoulos, A., Denning, D.W., Nierman, W.C. 2012. Sequencing of mitochondrial genomes of nine Aspergillus and Penicillium species identifies mobile introns and accessory genes as main sources of genome size variability. BMC Genomics, 13.

Wu, F., Bhatnagar, D., Bui-Klimke, T., Carbone, I., Hellmich, R., Munkvold, G., Paul, P., Payne, G., Takle, E. 2011. Climate change impacts on mycotoxin risks in US maize. World Mycotoxin J, 4: 79-93.

Reese, B. N., Payne, G. A., Nielsen, D. M. and Woloshuk, C. P. 2011. Gene Expression Profile and Response to Maize Kernels by Aspergillus flavus. Phytopathology, 101: 797-804.

Georgianna, D. R., Fedorova, N. D., Burroughs, J. L., Dolezal, A. L., Bok, J. W., Horowitz-Brown, S., Woloshuk, C.P., Yu, J., Keller, N.P., Payne, G.A. 2010. Beyond aflatoxin: four distinct expression patterns and functional roles associated with Aspergillus flavus secondary metabolism gene clusters. Mol Plant Pathol, 11: 213-226.

Eller, M. S., Payne, G. A. and Holland, J. B. 2010. Selection for reduced fusarium ear rot and fumonisin content in advanced backcross maize lines and their topcross hybrids. Crop Sci, 50: 2249-2260.

Chang, K. Y., Georgianna, D. R., Heber, S., Payne, G. A. and Muddiman, D. C. 2010. Detection of alternative splice variants at the proteome level in Aspergillus flavus. J Proteome Res, 9: 1209-1217.

Payne, G. A. and Yu, J.  2010.  Ecology, Development and Gene Regulation in Aspergillus flavus. Pages 157-171 in, Aspergillus: Molecular Biology and Genomics. Eds. Masayuki Machida and Katsuya Gomi.  Caister Academic Press, Norfolk, UK. 238 pages.

Nicholson, M. J., Koulman, A., Monahan, B. J., Pritchard, B. L., Payne, G. A. and Scott, B. 2009. Identification of two aflatrem biosynthesis gene loci in Aspergillus flavus and metabolic engineering of Penicillium paxilli to elucidate their function. Appl Environ Microb, 75: 7469-7481.

Georgianna, D. R. and Payne, G. A. 2009. Genetic regulation of aflatoxin biosynthesis: From gene to genome. Fungal Genet Biol, 46: 113-125.

Cleveland, T. E., Yu, J. J., Fedorova, N., Bhatnagar, D., Payne, G. A., Nierman, W. C., et al. 2009. Potential of Aspergillus flavus genomics for applications in biotechnology. Trends Biotechnol, 27: 151-157.

Yu, J. J., Payne, G. A., Nierman, W. C., Machida, M., Bennett, J. W., Campbell, B. C., et al. 2008. Aspergillus flavus genomics as a tool for studying the mechanism of aflatoxin formation. Food Addit Contam A, 25: 1152-1157.

Smith, C. A., Robertson, D., Yates, B., Nielsen, D. M., Brown, D., Dean, R. A., Payne, G.A. 2008. The effect of temperature on Natural Antisense Transcript NAT expression in Aspergillus flavus. Curr Genet, 54: 241-269.

Machida, M., Terabayashi, Y., Sano, M., Yamane, N., Tamano, K., Payne, G. A., et al. 2008. Genomics of industrial Aspergilli and comparison with toxigenic relatives. Food Addit Contam A, 25: 1147-1151.

Johnson, D. N., Egner, P. A., OBrian, G., Glassbrook, N., Roebuck, B. D., Sutter, T. R., Payne, G.A., Kensler, T.W., Groopman, J.D. 2008. Quantification of urinary aflatoxin B-1 dialdehyde metabolites formed by aflatoxin aldehyde reductase using isotope dilution tandem mass spectrometry. Chem Res Toxicol, 21: 752-760.

Holmes, R. A., Boston, R. S. and Payne, G. A. 2008. Diverse inhibitors of aflatoxin biosynthesis. Appl Microbiol Biot, 78: 559-572.

Georgianna, D. R., Hawkridge, A. M., Muddiman, D. C. and Payne, G. A. 2008. Temperature-dependent regulation of proteins in Aspergillus flavus: Whole organism stable isotope labeling by amino acids. J Proteome Res, 7: 2973-2979.

Eller, M. S., Robertson-Hoyt, L. A., Payne, G. A. and Holland, J. B. 2008. Grain yield and fusarium ear rot of maize hybrids developed from lines with varying levels of resistance. Maydica, 53: 231-237.

Eller, M., Holland, J. and Payne, G. 2008. Breeding for improved resistance to fumonisin contamination in maize. Toxin Rev, 27: 371-389.

Collier, T. S., Hawkridge, A. M., Georgianna, D. R., Payne, G. A. and Muddiman, D. C. 2008. Top-down identification and quantification of stable isotope labeled proteins from Aspergillus flavus using online nano-flow reversed-phase liquid chromatography coupled to a LTQ-FTICR mass spectrometer. Anal Chem, 80:4994-5001.

Bhatnagar, D., Rajasekaran, K., Payne, G. A., Brown, R. L., Yu, J. and Cleveland, T. E. 2008. The ‘omics’ tools: genomics, proteomics, metabolomics and their potential for solving the aflatoxin contamination problem. World Mycotoxin J, 1: 3-12.

Yu, J. J., Ronningy, C. M., Wilkinsony, J. R., Campbell, B. C., Payne, G. A., Bhatnagar, D., et al. 2007. Gene profiling for studying the mechanism of aflatoxin biosynthesis in Aspergillus flavus and A. parasiticus. Food Addit Contam A, 24: 1035-1042.

Smith, C. A., Woloshuk, C. P., Robertson, D. and Payne, G. A. 2007. Silencing of the aflatoxin gene cluster in a diploid strain of Aspergillus flavus is suppressed by ectopic aflR expression. Genetics, 176: 2077-2086.

Rokas, A., Payne, G., Fedorova, N. D., Baker, S. E., Machida, M., Yu, J., et al. 2007. What can comparative genomics tell us about species concepts in the genus Aspergillus? Stud Mycol, 11-17.

Robertson-Hoyt, L. A., Kleinschmidt, C. E., White, D. G., Payne, G. A., Maragos, C. M. and Holland, J. B. 2007. Relationships of resistance to Fusarium ear rot and fumonisin contamination with agronomic performance of maize. Crop Sci, 47: 1770-1778.

Robertson-Hoyt, L. A., Betran, J., Payne, G. A., White, D. G., Isakeit, T., Maragos, C. M., et al. 2007. Relationships among resistances to Fusarium and Aspergillus ear rots and contamination by fumonisin and aflatoxin in maize. Phytopathology, 97: 311-317.

OBrian, G. R., Georgianna, D. R., Wilkinson, J. R., Yu, J., Abbas, H. K., Bhatnagar, D., Cleveland, T.E., Nierman, W., Payne, G.A. 2007. The effect of elevated temperature on gene transcription and aflatoxin biosynthesis. Mycologia, 99: 232-239.

He, Z. M., Price, M. S., OBrian, G. R., Georgianna, D. R. and Payne, G. A. 2007. Improved protocols for functional analysis in the pathogenic fungus Aspergillus flavus. BMC Microbiol, 7.

Du, W., Obrian, G. R. and Payne, G. A. 2007. Function and regulation of aflJ in the accumulation of aflatoxin early pathway intermediate in Aspergillus flavus. Food Addit Contam, 24: 1043-1050.

Cary, J. W., OBrian, G. R., Nielsen, D. M., Nierman, W., Harris-Coward, P., Bhatnagar, D., Cleveland, T.E., Payne, G.A., Calvo, A.M. 2007. Elucidation of veA-dependent genes associated with aflatoxin and sclerotial production in Aspergillus flavus by functional genomics. Appl Microbiol Biot, 76: 1107-1118.

Robertson-Hoyt, L. A., Jines, M. P., Balint-Kurti, P. J., Kleinschmidt, C. E., White, D. G., Payne, G. A., et al. 2006. QTL mapping for fusarium ear rot and fumonisin contamination resistance in two maize populations. Crop Sci, 46: 1734-1743.

Robertson, L. A., Kleinschmidt, C. E., White, D. G., Payne, G. A., Maragos, C. M. and Holland, J. B. 2006. Heritabilities and correlations of fusarium ear rot resistance and fumonisin contamination resistance in two maize populations. Crop Sci, 46: 353-361.

Price, M. S., Yu, J. J., Nierman, W. C., Kim, H. S., Pritchard, B., Jacobus, C. A., Bhatnagar, D.,Cleveland, T.E., Payne, G.A. 2006. The aflatoxin pathway regulator AfIR induces gene transcription inside and outside of the aflatoxin biosynthetic cluster. Fems Microbiol Lett, 255: 275-279.

Johansson, A. S., Whitaker, T. B., Hagler, W. M., Bowman, D. T., Slate, A. B. and Payne, G. 2006. Predicting aflatoxin and fumonisin in shelled corn lots sing poor-quality grade components. J AOAC Int, 89: 433-440.

Payne, G. A., Nierman, W. C., Wortman, J. R., Pritchard, B. L., Brown, D., Dean, R. A., et al. 2006 Whole genome comparison of Aspergillus flavus and A-oryzae. Med Mycol, 44, S9-S11.

Xue, H. Q., Isleib, T. G., Payne, G. A., Novitzky, W. F. and Obrian, G. .2005. Aflatoxin production in peanut lines selected to represent a range of linoleic acid concentrations. J Food Protect, 68: 126-132.

Price, M. S., Shannon, B. C. B., Sabrina, T. B., Robert, A. K. B. and Payne, G. A. .2005. Aflatoxin conducive and non-conducive growth conditions reveal new gene associations with aflatoxin production. Fungal Genet Biol, 42: 506-518.

Yu, J. J., Chang, P. K., Ehrlich, K. C., Cary, J. W., Bhatnagar, D., Cleveland, T. E., Payne, G.A., Linz, J.E., Woloshuk, C.P., Bennett, J.W. 2004. Clustered pathway genes in aflatoxin biosynthesis. Appl Environ Microb, 70: 1253-1262.

Pirttila, A. M., McIntyre, L. M., Payne, G. A. and Woloshuk, C. P. 2004. Expression profile analysis of wild-type and fec1 mutant strains of Fusarium verticillioides during fumonisin biosynthesis. Fungal Genet Biol, 41: 647-656.

Moore, K. G., Price, M. S., Boston, R. S., Weissinger, A. K. and Payne, G. A. 2004. A chitinase from Tex6 maize kernels inhibits growth of Aspergillus flavus. Phytopathology, 94: 82-87.

Bush, B. J., Carson, M. L., Cubeta, M. A., Hagler, W. M. and Payne, G. A. 2004. Infection and fumonisin production by Fusarium verticillioides in developing maize kernels. Phytopathology, 94: 88-93.

Xue, H. Q., Isleib, T. G., Payne, G. A., Wilson, R. F., Novitzky, W. P. and O’Brian, G. 2003. Comparison of aflatoxin production in normal- and high-oleic backeross-derived peanut lines. Plant Dis, 87: 1360-1365.

Scheidegger, K. A. and Payne, G. A. 2003 Unlocking the secrets behind secondary metabolism: A review of Aspergillus flavus from pathogenicity to functional genomics. J Toxicol-Toxin Rev, 22: 423-459.

OBrian, G. R., Fakhoury, A. M. and Payne, G. A. 2003 Identification of genes differentially expressed during aflatoxin biosynthesis in Aspergillus flavus and Aspergillus parasiticus. Fungal Genet Biol, 39: 118-127.

Brown, R. L., Brown-Jenco, C. S., Bhatnagar, D. and Payne, G. A. 2003 Construction and preliminary evaluation of an Aspergillus flavus reporter gene construct as a potential tool for screening aflatoxin resistance. J Food Protect, 66: 1927-1931.

Price, M. S., Classen, J. J. and Payne, G. A. 2001 Aspergillus niger absorbs copper and zinc from swine wastewater. Bioresource Technol, 77: 41-49.

Nielsen, K., Payne, G. A. and Boston, R. S. 2001 Maize ribosome-inactivating protein inhibits normal development of Aspergillus nidulans and Aspergillus flavus. Mol Plant Microbe In, 14: 164-172.

Brown, R. L., Cleveland, T. E., Woloshuk, C. P., Payne, G. A. and Bhatnagar, D. 2001 Growth inhibition of a Fusarium verticillioides GUS strain in corn kernels of aflatoxin-resistant genotypes. Appl Microbiol Biot, 57: 708-711.

Reviews and Book Chapters

Bhatnagar, D., Payne, G., Klich, M. and Leslie, J. F. 2014 Identification of Toxigenic Aspergillus and Fusarium Species in the Maize Grain Chain. In: Mycotoxin Reduction in Grain Chains. John Wiley & Sons, Ltd, pp. 9-25.

Yu, J., Nierman, W. C., Bennett, J. W., Cleveland, T. E. Campbell, C. C., Bhatnagar, D., Dean, R. A., and Payne, G. A. 2010. Genetics and Genomics of Aspergillus flavus. In: Rai, M. K. and Kovics, G. (ed). Progress in Mycology. Springer Verlag. Chapter 2. Pages 51-73.

Payne, G. A. Genomics of Aspergillus flavus. 2007. In, The Aspergilli: Genomics, Medical Applications, Biotechnology, and Research Methods. Osmoni and Goldman CRC Press.

Payne, G. A. and Cubeta, M. A. 2005. Biology and detection of fungal pathogens of humans and plants. In Microbial Forensics, Eds. R.G. Breeze, B. Budowle, and S. E. Schutzer. Elsevier Academic Press. New York.

Robertson, L.A., Payne, G.A., and Holland, J.B. 2005. Marker Assisted Breeding for Host Resistance to Mycotoxin Contamination. Aflatoxin and Food Safety, edited by H. Abbas. CRC Press.

Price, M. S. and Payne, G. A. 2004. Genomics and gene regulation of the aflatoxin biosynthetic pathway. p.387-404. In Z. An (ed.), Handbook of Industrial Mycology. Marcel Dekker, Inc., New York.

Bhatnagar, D., G. A. Payne, T. E. Cleveland, and J. F. Robens. 2004. Mycotoxins: Current Issues in USA. Pages 17-47, in Meeting the Mycotoxin Menace, Eds. H. Barug, H. P. van Egmond, R. Lopez-Garcia, W. A. van Osenbruggen, and A. Visconti. Wageningen Academic Publishers, Netherlands.

Robertson, L.A., Payne, G.A., and Holland, J.B. 2005. Marker Assisted Breeding for Host Resistance to Mycotoxin Contamination. Aflatoxin and Food Safety, edited by H. Abbas. CRC Press.

Fakhoury, A. K. and Payne, G. A. 2003. Genomics of filamentous fungi: A general overview. In D. K. Arora, ed. Handbook of Fungal Biotechnology. 2nd Ed. Volume 20 of the Mycology Series.

Wu, F., Bhatnagar, D., Bui-Klimke, T., Carbone, I., Hellmich, R., Munkvold, G., Paul, P., Payne, G., Takle, E. 2011. Climate Change Impacts on Mycotoxin Risks in US Maize. World Mycotoxin Journal. 4: 79-93.

Richard, J. L. and G. A. Payne. 2003. Eds., Mycotoxins: Risks in Plant, Animal, and Human Systems. Task Force Report 139. Council for Agricultural Science and Technology. Ames, Iowa. 199 pages.