Paul Cullen excels in mentoring postdoctoral scholars

Paul Cullen, professor and director of graduate studies, is the recipient of the 2021-22 Distinguished Postdoctoral Mentor Award, which recognizes UB faculty members who excel in the mentoring of postdoctoral scholars. The award is bestowed on faculty members who not only teach their mentees, but also serve as an advocate, adviser and positive role model. Read the news story by Charles Anzalone.

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Cullen receives Distinguished Postdoc Mentor Award


Published February 3, 2023

“Dr. Cullen singlehandedly shaped my academic trajectory and is the main reason why I decided to stay in academia. ”
Beatriz Gonzalez, postdoctoral fellow
Department of Biological Sciences
Paul Cullen.

Paul Cullen

Paul Cullen, professor and director of graduate studies in the Department of Biological Sciences, is the recipient of the 2021-22 Distinguished Postdoctoral Mentor Award, which recognizes UB faculty members who excel in the mentoring of postdoctoral scholars.

The annual award, established in 2009, is presented by the Graduate School’s Office of Postdoctoral Scholars. It supports faculty members who not only teach their mentees, but also serve as an advocate, adviser and positive role model.

“The Office of Postdoctoral Scholars was thrilled at this year’s nomination pool,” says Kristen Ashare, director of the Office of Postdoctoral Scholars. “We are continuing to develop opportunities for highlighting mentorship and positive mentor-mentee relationship development here on campus. 

“This annual award honors important work that otherwise goes unseen,” Ashare says. “I’m so grateful to Dr. Cullen and his dedication to mentorship that is recognized with this award.”

Cullen’s research investigates how cells sense changes in the environment and make decisions. He received his PhD from Washington University in St. Louis, where he studied nitrogen sensing and signaling in bacteria under Robert Kranz, professor of biology. Cullen was a postdoctoral fellow at the University of Oregon with George Sprague, now professor emeritus of biology, where he explored glucose signaling in the regulation of filamentous growth in yeast, primarily by mucin sensors and Mitogen-Activated Protein Kinase (MAPK) pathways. His interest in microbial signal transduction has continued as a faculty member in the UB Department of Biological Sciences for the past 18 years.

“All cells sense and respond to extracellular signals,” Cullen explains. “Sensing and relaying changes in the extracellular milieu is mediated by signal transduction pathways. One type of evolutionary conserved signaling pathway are Mitogen-Activated Protein Kinase (MAPK) pathways that can be controlled by G-proteins like the ubiquitous Cdc42. Cdc42-driven MAPK pathways are found in many eukaryotes and function in diverse ways to regulate cell differentiation, cell cycle progression and the response to stress,” he says. “We found a mucin-type protein that regulates MAPK pathways and are interested in learning about what these sticky molecules are sensing and how they trigger changes in cell shape.

“An interesting feature of MAPK pathways is that they can share components with other pathways in the cell,” Cullen notes. “How pathways that function in integrated networks induce the ‘right’ response is not well understood. Furthermore, inappropriate regulation of MAPK pathways can lead to cross talk, which is an underlying cause of diseases including cancers, immune diseases, inflammation and neurodegenerative disorders.”

Cullen’s nomination for the Distinguished Postdoc Mentor Award included a letter of endorsement from Beatriz Gonzalez, postdoctoral fellow in the Department of Biological Sciences.

“Dr. Cullen singlehandedly shaped my academic trajectory and is the main reason why I decided to stay in academia,” Gonzalez wrote in her nomination letter.

“During my experience as a graduate student, I had other advisers who did not drive my attention to an academic career. I was considering non-academic career choices until I met Dr. Cullen during an international collaboration, and I had the opportunity to spend 4 months in his laboratory.”

Gonzalez called her experience with Cullen “really fruitful in terms of research, writing and thinking.” As supervisor, Cullen “completely changed my vision about research and science,” she said.

“Dr. Cullen is a positive role model who has reinvigorated my love of science,” she wrote. “He really respects and understands my goals and makes a real effort to help me to achieve my aspirations. He also drives passion toward his research and transmits this passion to me.

“Dr. Cullen puts my career goals as a priority. It is not uncommon to hear from my postdoctoral colleagues that they are just cogs on a greater machine, their primary goal being achieving the PI’s goals,” Gonzalez wrote. “Rather, I feel that I am working on my future in Dr. Cullen’s laboratory and exploring exciting research together with my colleagues.

“In addition, as a woman, we are uniquely vulnerable and face more obstacles that other people in our same situation. I was so lucky to have one mentor like Dr. Cullen, who helps and supports me to keep fighting and pursue my dreams and goals against the odds.”

Gonzalez’s letter “impressed” the selection committee, Ashare said.

“It truly takes a mentor going above and beyond to close these leaks in the talent pipeline and we are so glad that Dr. Beatriz Gonzalez shared her story,” she said.

Faculty Profile

  • Paul J. Cullen


    Paul J. Cullen.

    Paul J. Cullen


    Paul J. Cullen


    Director of Graduate Studies
    Distinguished Post Doc Mentor Award 2023
    Milton Plesur Excellence in Teaching Award
    UB Exceptional Scholar — Sustained Achievement Award

    Research Interests

    Signaling (MAPK) Pathways that Control Cell Polarity and Cell Differentiation


    • PhD, Washington University, Saint Louis
    • BA, University of Illinois

    Office Hours

    Research Summary

    The Cullen Lab is interested in signal transduction pathways. All cells sense and respond to changes in the environment. Sensing and relaying changes in the extracellular milieu is mediated by signal transduction pathways. One type of evolutionary conserved signaling pathway are Mitogen-Activated Protein Kinase (MAPK) pathways. MAPK pathways are found in all eukaryotes and function in diverse ways to regulate cell differentiation, cell cycle progression, and the response to stress. An interesting feature of MAPK pathways is that they can share components with other pathways in the cell. How pathways that function in integrated networks induce the ‘right’ response is not well understood. Furthermore, inappropriate regulation of MAPK pathways can lead to improper signaling, which is an underlying cause of diseases including cancers, immune diseases, inflammation, and neurodegenerative disorders.

    Our laboratory uses the model organism budding yeast to study MAPK pathways. In yeast, MAPK pathways control the response to stress and orchestrate cell differentiation to specialized cell types. Using this model, we have been able to identify new regulators of MAPK pathways and new mechanisms for their regulation. For example, we showed that mucins that regulate MAPK pathways in yeast undergo processing and release of an inhibitory extracellular glycodomain. More recently, we have identified an adaptor that regulates and might help to insulate the pathway (the pathway we study shares components with other MAPK pathways). We are currently working on how positional cues impact MAPK pathway signaling through the Rho GTPase Cdc42. 

    The specific MAPK that we study in our laboratory regulates a cell differentiation response called filamentous growth. Filamentous growth is a fungal-specific growth mode that occurs in yeast and many other fungal species. During filamentous growth, cells change their shape and grow as branched filaments. In some species of pathogenic microorganisms, filamentous growth is required for virulence. Among the signaling pathways that regulate filamentous growth is an ERK-type MAPK pathway called the filamentous growth (fMAPK) pathway. Studying how MAPK pathways regulate filamentous growth in yeast can shine light on MAPK-dependent differentiation responses in general as well as the molecular basis of fungal pathogenesis.

    The proteins that regulate the fMAPK pathway in yeast have been identified by our laboratory and other laboratories. A signalling mucin-type glycoprotein (Msb2), together with polar landmarks, regulates the cell’s major polarity GTPase Cdc42p, which activates a canonical p21-MAPKKK-MAPKK-MAPK kinase cascade. Key questions surrounding the regulation of the fMAPK pathway remain unaddressed. One question is how does the mucin Msb2 regulate the GTPase module? A second question is how is a specific signal sent by a pathway that shares components with other MAPK pathways in the cell? Our lab is approaching these questions in different ways:

    1. By the development and utilization of genomic and proteomic approaches, we have discovered and are continuing to identify proteins that regulate the filamentous growth pathway. For example, we used the fact that the extracellular domain of Msb2 is shed to develop a high throughput screening approach called secretion profiling to identify new regulators of Msb2/fMAPK.

    2. By building new assays to measure and quantitate filamentous growth, we are learning more about the molecular basis of the response. We are currently utilizing genetics, microscopy, and bioinformatics analysis to define aspects of filamentous growth regulation.

    3. Biochemistry, molecular biology, and in vivo protein-interaction approaches (e.g. two-hybrid analysis and FRET) are being employed, which are critical to determine how proteins that regulate MAPK pathways function to induce a response.

    Selected Publications

    • Pujari, A.N. and Cullen, P.J. (2024) Modulators of MAPK pathway activity during filamentous growth in Saccharomyces cerevisiae G3 (Bethesda):jkae072. doi: 10.1093/g3journal/jkae072.
    • Vandermeulen, M.D. and Cullen, P.J. (2023) Ecological inducers of the yeast filamentous growth MAPK pathways reveal environment-dependent roles for pathway components. mSPHERE (5):e0028423. doi: 10.1128/msphere.00284-23. 
    • Gonzalez, B., Mirzaei, M., Basu, S., Pujari, A. Vandermeulen, M.D., Prabhakar, A., Cullen, P.J. (2023) New Features Surrounding the Cdc42-Ste20 Module that Regulates MAP Kinase Signaling in Yeast. Journal of Biological Chemistry 299 (11):105297. doi: 10.1016/j.jbc.2023.105297.
    • Vijjamarri, A.K., Niu, X., Vandermeulen, M.D., Onu, C., Zhang, F., Qiu, H., Gupta, N., Gaikwad, S., Greenberg, M.L., Cullen, P.J., Lin, Z., Hinnebusch, A.G. (2023) Decapping factor Dcp2 controls mRNA abundance and translation to adjust metabolism and filamentation to nutrient availability. eLIFE DOI: 10.7554/eLife.85545
    • González, B., Aldea, M. and Cullen, P.J. (2023) Chaperone-Dependent Degradation of Cdc42 Promotes Cell Polarity and Shields the Protein from Aggregation, Molecular and Cellular Biology doi: 10.1080/10985549.2023.2198171.
    • Gonzalez, B. and Cullen, P.J. (2022) Cdc42 turnover impacts the filamentation MAPK pathway in yeast  Journal of Cell Biology 221( 12):e202112100. doi: 10.1083/jcb.202112100.
    • Vandermeulen, M.D. and Cullen, P.J. (2022) Gene by Environment Interactions Reveal New Regulatory Aspects of Signaling Network Plasticity  PLoS Genetics V18(1).
    • Prabhakar, A., Gonzalez, B., Dionne, H., Basu, S. and Cullen, P.J. Spatiotemporal Control of Pathway Sensors and Cross-Pathway Feedback Regulate a Cell Differentiation MAPK Pathway in Yeast (2021) Journal of Cell Science Jul 9:jcs.258341. doi: 10.1242/jcs.258341. PMID: 34240741.
    • The signaling network of the yeast NADPH oxidase Yno1p (2021) Weber, M. Basu, S., Gonzalez, B., Greslehner, G., Singer, S., Haskova, D., Hasek, J., Breitenbach, M., Gourlay, C., Cullen, P.J. *, and Rinnerthaler, M. *Antioxidants (Basel) 10(2):322. doi: 10.3390/antiox10020322
    • Jamalzadeh, S. Pujari, A.N. and Cullen, P.J. (2020) A Rab Escort Protein Regulates the MAPK Pathway That Controls Filamentous Growth in Yeast, Scientific Reports. 10(1) : 22184. doi: 10.1038/s41598-020-78470-4.
    • Slow growth and increased spontaneous mutation frequency in respiratory deficient yeast suppressed by a dominant mutation in ATP3. (2020) Li, J., Rinnerthaler, M., Hartl, J., Weber, M., Karl, T., Breitenbach-Koller, H., Mülleder, M., Vowinckel, J., Mattanovich, D., Sauer, M., Marx, H., Greslehner, G., Geltinger, F., Burhans, B., Grant, C., Doronina, V., Ralser, M., Streubel, K., Grabner, C., Jarolim, S., Moßhammer, C., Gourlay, C.W., Cullen, P.J. Liti, G., Ralser, M., and Breitenbach, M. G3: Genes Genomes Genetics. 10(12):4637-4648. doi: 10.1534/g3.120.401537.
    • New Aspects of Invasive Growth Regulation Identified by Functional Profiling of MAPK Pathway Targets in Saccharomyces cerevisiae  Vandermeulen, M.D. and Cullen, P.J. (2020) GENETICS doi: 10.1534/genetics.120.303369
    • Sho1p Connects Glycolysis to Ras1-cAMP Signaling and is Required for Microcolony Formation in Candida albicans" (2020) Rohitashw Kumar, Malabika Maulik, Ruvini Pathirana, Cullen, P.J. and M. Edgerton mSPHERE 5(4):e00366-20. doi: 10.1128/mSphere.00366-20.
    • Binding and release of Fe(III) complexes from glucan particles for delivery of T1 MRI contrast agents (2020) Patel, A., Asik, D. Snyder, E.M., Delillo, A.E., Cullen, P.J., and Morrow, J.R. ChemMedChem 15(12):1050-1057. 
    • Prabhakar, A., Chow, J., Siegel, A.J., and Cullen, P.J. (2020) Regulation of Intrinsic Polarity by a MAPK Pathway  Journal of Cell Science pii: jcs.241513. doi: 10.1242/jcs.241513.
    • Basu, S., González, B., Li, B., Kimble, G., Kozminski, K.G. and Cullen, P.J. (2020) Functions for Cdc42p BEM Adaptors in Regulating a Differentiation-Type MAP Kinase Pathway. Molecular Biology of the Cell Jan 15:mbcE19080441. doi: 10.1091/mbc.E19-08-0441.
    • Prabhakar, A., Vadaie, N., Krzystek, T., and Cullen, P.J. (2019) Proteins That Interact with the Mucin-Type Glycoprotein Msb2p Include a Regulator of the Actin Cytoskeleton Biochemistry 58 (48):4842-4856. doi: 10.1021/acs.biochem.9b00725.
    • Patel, A., Asik, D., Cullen, P.J. and Morrow, J.R.  (2019) MRI and fluorescence studies of Saccharomyces cerevisiae loaded with a bimodal Fe(III) T1 contrast agent. Journal of Inorganic Biochemistry. 201:110832. doi: 10.1016/j.jinorgbio.2019.110832.
    • McCall, A.D., Pathirana, R.U., Prabhakar, A., Cullen, P.J., and Edgerton, M. (2019) Candida albicans biofilm development is governed by cooperative attachment and adhesion maintenance proteins. Nature Publishing Journal Biofilms 5:21. doi: 10.1038/s41522-019-0094-5.
    • Chow J., Starr, I., Jamalzadeh, S., Muniz, O., Kumar, A., Gokcumen, O., Ferkey, D.M. and Cullen, P.J. (2019) Filamentation Regulatory Pathways Control Adhesion-Dependent Surface Responses in Yeast. Genetics.  212(3):667-690. Article.
    • Colloidal Gels with Tunable Mechanomorphology Regulate Endothelial Morphogenesis (2019) Nair, S.K., Basu, S., Sen, B., Lin, M.S., Kumar, A.N., Yuan, Y., Cullen, P.J., Sarkar, D. Scientific Reports  9(1):1072.
    • Chow, J., Dionne, H.M, Prabhakar, A., Mehrotra, A., Somboonthum, J., Gonzalez, B., Edgerton, M., and Cullen, P.J. (2019) Aggregate Filamentous Growth Responses in Yeast mSPHERE V4(2) e00702-18.
    • Breitenbach, M., Rinnerthaler, M., Weber, M., Breitenbach-Koller, H., Karl, T. Cullen, P.J., Basu, S., Haskova, D., and Hasek, J. (2018) The defense and signaling role of NADPH oxidases in eukaryotic cells. Wien. Med. Wochenschr. 168: 286-299.
    • Sehgal, N.,  Sylves, M.E., Sahoo, A., Chow, J., Walker, S., Cullen, P.J., and Berry, J.O. (2018) CRISPR gene editing in yeast: An experimental protocol for an upper-division undergraduate laboratory course.Biochemistry and Molecular Biology Education doi: 10.1002/bmb.21175.
    • Chow, J., Notaro, M., Prabhakar, A., Free, S.J., and Cullen, P.J. (2018) Impact of Fungal MAPK Pathway Targets on the Cell Wall Journal of Fungi  V4:93.
    • Norman, K.L, Shively, C.A., De La Rocha, A.J., Basu, S., Cullen, P.J., and Kumar, A. (2018) Inositol Polyphosphates Regulate and Predict Yeast Pseudohyphal Growth Phenotypes PLoS Genetics 14(6):e1007493.
    • González, B., Vázquez, J., Cullen, P.J, Mas, A., Beltran, G., and Torija, M.J. (2018) Aromatic amino acid-derived compounds induce morphological changes and modulate the cell growth of wine yeast species. Frontiers of Microbiology  9:670.
    • Kumar, R., Breindel, C., Saraswat, D., Cullen, P.J., and M. Edgerton (2017) Candida albicans Sap6 amyloid regions function in cellular aggregation and zinc binding, and contribute to zinc acquisition. Scientific Reports 7:1p2908.
    • Gonzalez, B. Mas, A., Beltran, G., Cullen, P.J., Torija, M.J. (2017) Retrograde Mitochondria Pathway Regulates Ethanol-Inducible Filamentous Growth in Yeast Frontiers in Physiology 8:148. doi: 10.3389/fphys.2017.00148.
    • Woolford, C.A., Lagree, K., Xu, W., Aleynikova, T., Adhikari, H., Sanchez, H., Cullen, P.J., Lanni, F., Andes, D.R., and AP Mitchell (2016) Biofilm formation in the absence of Candida albicans biofilm transcriptional activators. PLoS Genetics. DOI:10.1371/journal.pgen.1006487.
    • Basu, S., Vadaie, N., Pradhakbar, A. Li, B., Pitoniak, A., Adhikari, H., Chavel, C., and Cullen, P.J. (2016) Spatial Landmarks Regulate a Cdc42p-Dependent MAPK Pathway to Control Differentiation and the Response to Positional Compromise. PNAS. 113(14):E2019-28.
    • Saraswat, D. Kumar, R., Pande, T. Edgerton M. and Cullen, P.J. (2016) Signaling Mucin Msb2 Regulates Adaptation to Thermal Stress in Candida albicans. Mol. Microbiology100(3):425-41.
    • Cullen, P.J. and Edgerton M. (2016) Unmasking Fungal Pathogens by Studying MAPK-Dependent Cell Wall Regulation in Candida albicans. Virulence. Apr 18:1-4.
    • Cullen, P.J. (2015) The Plate-Washing Assay: A Simple Test for Filamentous Growth in Budding Yeast. Cold Spring Harbor Laboratory Protocols. (2):168-71.
    • Cullen, P.J. (2015) Evaluating Yeast Filamentous Growth at the Single-Cell Level. Cold Spring Harbor Laboratory Protocols. (3): 272-5.
    • Cullen, P.J. (2015) Evaluating the Activity of the Filamentous Growth Mitogen Activated Protein Kinase Pathway in Yeast. Cold Spring Harbor Laboratory Protocols. (3):276-83.
    • Cullen, P.J. (2015) Biofilm/Mat Assays for Budding Yeast. Cold Spring Harbor Laboratory Protocols. (2): 172-5.
    • Cullen, P.J. (2015) Investigating Filamentous Growth and Biofilm/Mat Formation in Budding Yeast. Cold Spring Harbor Laboratory Protocols. (3): 235-8.
    • Adhikari, H., Caccamise, L.M., Pande, T., and Cullen, P.J (2015) Comparative Analysis of Transmembrane Regulators of the Filamentous Growth MAPK Pathway Uncovers Functional and Regulatory Differences. Eukaryotic Cell. 14(9):868-883.
    • Adhikari, H. Vadaie, N., Chow, J., Caccamise, L.M., Chavel, C.A., Li, B., Bowitch, A. Stefan, C.J., and Cullen, P.J. (2015) Role of the Unfolded Protein Response in Regulating the Mucin-Dependent Filamentous-Growth Mitogen Activated Protein Kinase Pathway. Molecular and Cellular Biology. 35(8):1414-32.
    • Adhikari, H. and Cullen, P.J. (2015) Role of Phosphatidylinositol Phosphate Signaling In the Regulation of the Filamentous Growth MAPK Pathway. Eukaryotic Cell. 14(4): 427-40.
    • Li, R. Puri, S. Tati, S. Cullen, P.J. and Edgerton, M. (2015) Candida albicans Cek1 MAPK signaling enhances fungicidal activity of salivary Histatin. Antimicrobial Agents and Chemotherapy. 59(6):3460-8.
    • Pitoniak A., Chavel C.A., Chow J., Smith J., Camara D., Karunanithi S., Li B., Wolfe K., Cullen P.J. (2015) Cdc42p-Interacting Protein Bem4p Regulates the Filamentous Growth MAP Kinase Pathway. Mol. Cell. Biol. Nov 10. pii: MCB.00850-14.
    • Chavel C.A., Caccamise L.M., Li B., Cullen P.J. (2014) Global regulation of a differentiation MAPK pathway in yeast. Genetics. V198(3):1309-28.
    • Adhikari, H. and Cullen P.J. (2014) Metabolic respiration induces AMPK- and Ire1p-dependent activation of the p38-Type HOG MAPK pathway. PLoS Genetics. V10(10):e1004734. doi: 10.1371/journal.pgen.1004734.
    • Puri, S., Kumar, R., Chadha, S., Tati S., Conti, H.R., Hube, B., Cullen, P.J, Edgerton M. (2012) Secreted Aspartic Protease Cleavage of Candida albicans Msb2 Activates Cek1 MAPK Signaling Affecting Biofilm Formation and Oropharyngeal Candidiasis. PLoS ONE. 7(11):e46020.
    • Karunanithi S., and Cullen P.J. (2012) The Filamentous Growth MAPK Pathway Responds to Glucose Starvation Through the Mig1/2 Transcriptional Repressors in Saccharomyces cerevisiaeGenetics. 192(3):869-87.
    • Meem M.H. and Cullen P.J. (2012) The impact of protein glycosylation on Flo11-dependent adherence in Saccharomyces cerevisiaeFEMS Yeast Research. 12(7):809-18.
    • Karunanithi S, Joshi J, Chavel C, Birkaya B, Grell L, and P.J. Cullen (2012)Regulation of Mat Responses by a Differentiation MAPK Pathway in Saccharomyces cerevisiae. PLoS ONE. 7(4): e32294
    • Cullen, P.J. and G.F. Sprague (2012) The regulation of filamentous growth in yeast. Genetics. V190:23-49.
    • Cullen, P.J. (2011) Post-translational Regulation of Signaling Mucins. Current Opinion of Structural Biology. V21 p590-596.
    • Karunanithi, S., Vadaie, N., Chavel, C., Birkaya, B., Joshi, J., Grell, L, and P.J. Cullen (2010) Shedding of the Mucin-like Flocculin Flo11p Reveals a New Aspect of Fungal Adhesion Regulation. Current Biology. 20:1-7.
    • Chavel, C.A., Dionne, H.S., Birkaya, B. Joshi, J., and P.J. Cullen (2010) New Regulators of a Differentiation MAPK Pathway. PLoS Genetics. 19; 6(3):e1000883
    • Pitoniak, A., Birkaya, B., Dionne H.M., Vadaie, N., and P. J. Cullen (2009) The Signaling Mucins Msb2 and Hkr1 Differentially Regulate the Filamentation MAPK Pathway and Contribute to a Multimodal Response. Molecular Biology of the Cell. V20 p3101-3114.
    • Birkaya, B. Maddi, A., Joshi, J., Free, S.J., and P.J. Cullen (2009) The Role of the Cell Wall Integrity Pathway and Cdc42-Dependent MAPK Pathway in Cell Wall Remodeling During Filamentous Growth. Eukaryotic Cell. V8 p1118-1133.
    • Abdullah, U. and P.J. Cullen (2009) The tRNA Modification Complex Elongator Regulates the Cdc42-Dependent MAPK Pathway that Controls Filamentous Growth In Yeast. Eukaryotic Cell. V8 p1362-1372.
    • Vadaie, N., Dionne, H., Nickerson, S.R., Akajagbor, D.S., Krysan, D.J., and Cullen, P.J. (2008) Cleavage of the signaling mucin Msb2 by the aspartyl protease Yps1 is required for MAPK activation in yeast. J. Cell Biology. 181: 1073-1081.
    • Cullen, P.J. (2007) Signaling Mucins: The New Kids on the MAPK Block. Critical Reviews in Eukaryotic Gene Expression
    • P.J. Cullen, Xu-Friedman, R., Delrow, J., and G. F. Sprague, Jr. (2006).Genome-wide analysis of a protein glycosylation deficiency. FEMS Yeast Research.
    • Devit, M.J., Cullen, P.J. Branson, M., Sprague, G.F., and Fields, S. (2005).A novel genetic screening method based on artificially forced protein interactions. Genome Research. 15 (4):560-565.
    • Cullen, P.J., Sabbagh Jr., W., Graham, E., Irick, M. van Olden, E.K., Neal, C., Delrow, J., Bardwell, L., and George F. Sprague, Jr. (2004). A signaling mucin at the head of the Cdc42- and MAPK-filamentous growth pathway in yeast.Genes & Development. 18:1695-1708.
    • Smith, G.R., Given, S.A., Cullen, P.J., and Sprague, G.F. (2002)Identification and Characterization of GTPase Activating Proteins (GAPs) for Cdc42. Eukaryotic Cell. V1 p469-480. 
    • Cullen, P.J., and Sprague, G.F., Jr. (2002) The Glc7p-interacting protein Bud14p attenuates pheromone response, filamentous growth, and polarized growth in Saccharomyces cerevisiaeEukaryotic Cell. V1:884-894.
    • Cullen, P.J., and Sprague, G.F., Jr. (2002)The roles of bud-site-selection proteins in haploid invasive growth in yeast. Mol. Biol. Cell. 13:2990-3004.
    • Cullen, P.J. and Sprague, G.F. (2000) Glucose depletion causes haploid invasive growth in yeast. PNAS. 97: 13619-13624.*See Commentary on this article: Madhani, H.D. (2000) Interplay of intrinsic and extrinsic signals in yeast differentiation. PNAS. 97: 13461-13463.
    • Cullen, P.J., Schultz, J., Horecka, J., Stevenson, B., and Jigami, Y., and Sprague, G.F. (2000) Defects in protein glycosylation cause SHO1-dependent activation of a STE12 signaling pathway in yeast. Genetics. 155: 1005-1018.