John E. Hayes, Ph.D.

John E. Hayes, Ph.D.

  • Professor of Food Science
  • Director, Sensory Evaluation Center
220 Rodney A. Erickson Food Science Building
University Park, PA 16802

Areas of Expertise

  • Sensory science
  • Human psychophysics
  • Taste biology
  • Drug delivery systems
  • Food choice
  • Ingestive behavior
  • Consumer acceptability


  • Postdoc, Brown University, Alcohol/Behavioral Genetics
  • Ph.D., University of Connecticut, Nutrition
  • Graduate Certificate, University of Connecticut, Quantitative Research Methods (Psychology)
  • M.S., Cornell University, Food Science (Sensory Science)
  • B.S., Cornell University, Food Science

Research Interests

Dr. Hayes leads a transdisciplinary research program that applies Sensory & Consumer Science to a broad range of topics, including ingestive behavior, flavor perception, and individual differences, as well as user centered drug delivery optimization, eCig flavor and use, and chemosensory changes due to COVID-19. Much of his work studies Food Choice within a biobehavioral framework by integrated traditional psychophysical and sensory methods with behavioral genetics to understand biological factors that may cause individuals to like and eat some foods but not others. During the COVID19 pandemic lockdown in Spring 2020, he initiated a new line of research on chemosensory loss and recovery following infection with SARS-CoV-2.

Recent projects in his laboratory have focused on:

  • Psychophysics of taste and flavor perception, especially bitterness, burn and astringency
  • Smell Loss & Recovery following SARS-CoV-2 infection
  • Rapid smell tests as a low cost screening tool for COVID19
  • Influences of genetic variation on food sensations, liking and intake
  • Sweet liking phenotype as a predictor of adiposity
  • Influence of personality on spicy food liking and intake
  • Perception and liking of non-nutritive sweeteners, including the genetic basis of individual differences across people
  • Factors which influence alcohol intake
  • Development of new psychophysical methods to predict consumer acceptability
  • Chemesthesis and bittern of oral pharmaceuticals
  • User acceptability of semisolid drug delivery systems for HIV prevention


  • 2021 – Sensory and Consumer Sciences Achievement Award (Institute of Food Technologists) 
  • 2016 – Barry Jacobs Memorial Award for Research on the Psychophysics of Human Taste and Smell (Association for Chemoreception Sciences)
  • 2015 – Food Quality and Preference Investigator of the Future Award (Pangborn Symposium / Elsevier)
  • 2013 – Roy C. Buck Award (Penn State College of Agricultural Sciences)
  • 2011 – Ajinomoto Award for Young Investigators in Gustation (Association for Chemoreception Sciences) 
  • 2006 – Rose Marie Pangborn Doctoral Scholarship (Pangborn Sensory Science Scholarship Fund)


Graduate Program in Neuroscience, Huck Institutes of the Life Sciences, Penn State

Graduate Program in Nutrition, College of Health and Human Development, Penn State

Peer Reviewed Publications

(also see


Iatridi V, Quadt L, Hayes JE, Garfinkel SN, Yeomans MR. 2021. Female sweet-likers have enhanced cross-modal interoceptive abilities. Appetite:105290.

Bakke AJ, Zaveri T, Higgins MJ, Ziegler GR, Hayes JE. 2021. Design aspects of vaginal applicators that influence acceptance among target users. Scientific Reports.

Brodock JL, Hayes JE, Masterson TD, Hopfer H. Differences in preferred fat level, sweetener type, and amount of added sugar in chocolate milk in a choice task relate to physical activity and orthorexia. Appetite. Accepted 13 March 2021.

Baker AN, Wilson SJ, Hayes JE. Flavor and product messaging are the two most important drivers of electronic cigarette selection in a choice-based task. Scientific Reports. 11: 4689.

Cunningham PM, Roe L, Hayes JE, Hetherington MM, Keller KL, & Rolls BJ. 2021. Development and Validation of the Reasons Individuals Stop Eating Questionnaire (RISE-Q): a novel tool to characterize satiation. Appetite. 161: 105127. 1 June 2021. doi: 10.1016/j.appet.2021.105127

Ferrante MJ, Moding KJ, Bellows LL, Bakke AJ, Hayes JE, Johnson SL. 2021. Examining front-of-package product names and ingredient lists of infant and toddler foods containing vegetables. Journal of Nutrition Education and Behavior. 53(2):96-102. ePub 1 Feb 2021. doi: 10.1016/j.jneb.2020.11.019


Gerkin RC, Ohla K, Veldhuizen MG, Joseph PV, Kelly CE, Bakke AJ, Steele KE, Farruggia MC, Pellegrino R, Pepino MY, Bouysset C, Soler GM, Pereda-Loth V, Dibattista M, Cooper KW, Croijmans I, Di Pizio A, Ozdener MH, Fjaeldstad AW, Lin C, Sandell MA, Singh PB, Brindha VE, Olsson SB, Saraiva LR, Ahuja G, Alwashahi MK, Bhutani S, D’Errico A, Fornazieri MA, Golebiowski J, Hwang LD, Öztürk L, Roura E, Spinelli S, Whitcroft KL, Faraji F, Fischmeister F, Heinbockel T, Hsieh JW, Huart C, Konstantinidis I, Menini A, Morini G, Olofsson JK, Philpott CM, Pierron D, Shields VDC, Voznessenskaya VV, Albayay J, Altundag A, Bensafi M, Bock MA, Calcinoni O, Fredborg W, Laudamiel C, Lim J, Lundström JN, Macchi A, Meyer P, Moein ST, Santamaría E, Sengupta D, Rohlfs-Dominguez P, Yanik H, GCCR Group Author, Hummel T, Hayes JE, Reed DR, Niv MY, Munger SD, Parma V. 2021. Recent smell loss is the best predictor of COVID-19 among individuals with recent respiratory symptoms. Chemical Senses. 46: bjaa081. doi: 10.1093/chemse/bjaa081

Trumbo PR, Appleton KM, de Graaf K, Hayes JE, Baer DJ, Beauchamp GK, Dwyer JT, Fernstrom JD, Klurfeld DM, Mattes RD, Wise PM. 2020. Measuring Sweetness in Foods, Beverages, and Diets: Toward Understanding the Role of Sweetness in Health. Advances in Nutrition. In press. ePub 3 December 2020. doi: 10.1093/advances/nmaa151

Pierron D, Pereda-Loth V, Mantel M, Moranges M, Bignon E, Alva O, Kabous J, Heiske M, Pacalon J, David R, Dinnella C, Spinelli S, Monteleone E, Farruggia MC, Cooper K, Sell E, Thomas-Danguin T, Bakke AJ, Parma V, Hayes JE, Letellier T, Ferdenzi C, Golebiowski J, Moustafa Bensafi M. 2020. Smell and taste changes are early indicators of the COVID-19 pandemic and political decision effectiveness. Nature Communications. 11:5152.

Baker AN, Bakke AJ, Branstetter SA, Hayes J. Harsh and sweet sensations predict acute liking of electronic cigarettes, but flavor does not affect acute nicotine intake: a pilot laboratory study in men. Nicotine & Tobacco Research. ePUB 12 October 2020. doi: 10.1093/ntr/ntaa209

Iatridi V, Armitage RM, Yeomans MR, Hayes JE. 2020. Effects of Sweet-Liking on Body Composition Depend on Age and Lifestyle: A Challenge to the Simple Sweet-Liking—Obesity Hypothesis. Nutrients. 12(9): 2702. doi: 10.3390/nu12092702

Gaby JM, Bakke AJ, Baker AN, Hopfer H, Hayes JE. 2020. Individual Differences in Thresholds and Consumer Preferences for Rotundone Added to Red Wine. Nutrients. 12(9): 2522. doi: 10.3390/nu12092552

Higgins MJ, Gipple JT, Hayes JE. 2021. Common bitter stimuli show differences in their temporal profiles before and after swallowing. Food Quality and Preference. 87: 104041.

Parma V, Ohla K, Veldhuizen MG, Niv MY, Kelly CE, Bakke AJ, Cooper KW, Bouysset C, Pirastu N, Dibattista M et al…. Hayes JE. 2020. More than smell – COVID-19 is associated with severe impairment of smell, taste, and chemesthesis. Chemical Senses.  doi: 10.1093/chemse/bjaa041

Lin C, Colquitt L, Wise P, Breslin P, Rawson NE, Genovese F, Maina I, Joseph P, Fomuso L, Slade L, Brooks D, Miclo A, Hayes JE, Sullo A, Reed DR. Studies of human twins reveal genetic variation that affects dietary fat perception. Chemical Senses. 45(6): 467-481. doi: 10.1101/chemse/bjaa036

Higgins MJ, Bakke AJ, Hayes JE. 2020. Personality traits and bitterness perception influence the liking and intake of pale ale style beers. Food Quality and Preference. 86: 103994.

Higgins MJ, Hayes JE. 2020. Discrimination of Isointense Bitter Stimuli in a Beer Model System. Nutrients. 12(6):1560.

Nolden AA, McGeary JE, Hayes JE. 2020. Predominant qualities evoked by quinine, sucrose, and capsaicin associate with PROP bitterness, but not TAS2R38 genotype. Chemical Senses. May 29;45(5):383-90.

Duffy VB, Hayes JE, Sharafi M. 2020. Interactions between retronasal olfaction and taste influence vegetable liking and consumption: a psychophysical investigation. Journal of Agriculture and Food Research. 2:100044.

Bakke AJ, Carney EM, Higgins MJ, Moding K, Johnson SL, Hayes JE. 2020. Blending dark green vegetables with fruits in commercially available infant foods makes them taste like fruit. Appetite. 150:104652. doi: 10.1016/j.appet.2020.104652

Etter NM, Breen SP, Alcala MIM, Ziegler GR, Hayes JE. 2020. Assessment of Midline Lingual Point-Pressure Somatosensation Using Von Frey Hair Monofilaments. Journal of Visualized Experiments: JoVE. 156. doi: 10.3791/60656.

    Gallo S, Byham-Gray L, Duffy VB, Hoffman HJ, Hayes JE, Rawal S. 2020. Associations of Olfactory Dysfunction with Anthropometric and Cardiometabolic Measures: Findings from the 2013-2014 National Health and Nutrition Examination Survey (NHANES). Physiology & Behavior. 112702. 


    Moding KJ, Ferrante MJ, Bellows LL, Bakke AJ, Hayes JE, Johnson SL. 2019. Nutritional Content and Ingredients of Commercial Infant and Toddler Food Pouches Compared to Other Packages Available in the United States. Nutrition Today. 54(6):305–312.

    Higgins MJ, Hayes JE. 2019. Regional variation of bitter taste and aftertaste in humans. Chem Senses. 44(9): 721–732.

    Gaby JM, Baker AN, Gravina S, Hayes JE. 2019. Vanillin modifies affective responses to but not burning sensations from ethanol in mixtures. Physiology & Behavior. 211: 112668. 

    Reyes MR, Gravina S, Hayes JE. 2019. Evaluation of sweetener synergy in humans by isobole analyses. Chem Senses. 44(8) 571–582. 

    Santagiuliana M, Sampedro Marigómez I, Broers L, Hayes JE, Piqueras-Fiszman B, Scholten E, Stieger M. 2019. Exploring variability in detection thresholds of microparticles through participant characteristics. Food Funct. 10: 5386-5397. 

    Nolden AA, Lenart G, Hayes JE. 2019. Putting out the fire – efficacy of common beverages in reducing oral burn from capsaicin. Physio Behav. 208:112557. 

    Fritts JR, Bermudez MA, Hargrove RL, Alla L, Fort C, Liang Q, Cravener TL, Rolls BJ, D'Adamo CR, Hayes JE, Keller KL. 2019. Using Herbs and Spices to Increase Vegetable Intake Among Rural Adolescents. Journal of Nutrition Education and Behavior

    Wang G, Bakke AB, Hayes JE, Hopfer H. 2019. Demonstrating cross-modal enhancement in a real food with a modified ABX test. Food Qual Pref

    Breen S, Etter NM, Ziegler GR, Hayes JE. 2019. Oral somatosensatory acuity is related to particle size perception in chocolate. Scientific Reports. 9(1):7437. 

    Perry DM, Byrnes NK, Heymann H, Hayes JE. 2019. Rejection of labrusca-type aromas in wine differs by wine expertise and geographic region. Food Qual Pref. 74: 147-154 -365. 

    Iatridi V, Hayes JE, Yeomans MR. 2019. Quantifying sweet taste liker phenotypes: time for some consistency in the classification criteria. Nutrients. 11(1):129-153. 

    Bakke AJ, Zaveri T, Ziegler GR, Hayes JE. 2019. Using sensory and consumer science in drug delivery system optimization: mixed methods in women of color as a case study. Food Qual Pref. 73: 293-302. 

    Iatridi V, Hayes JE, Yeomans MR. 2019. Reconsidering the classification of sweet taste liker phenotypes: a methodological review. Food Qual Pref. 72: 56-76. 

    Higgins MJ, Hayes JE. 2019. Learned color taste associations in a repeated brief exposure paradigm. Food Qual Pref. 71: 354-365.


    Zaveri T, Powell KA, Guthrie KM, Bakke AJ, Ziegler GR, Hayes JE. 2018. Qualitative exploration of intrinsic and extrinsic factors that influence acceptability of semisoft vaginal suppositories. BMC Women's Health. 18:170. 

    Wang G, Hayes JE, Ziegler GR, Roberts RF, Hopfer H. 2018. Dose-response relationships for vanilla flavor and sucrose in skim milk: evidence of synergy. Beverages. 4(4):73.

    Baker AN, Miranda AM, Garneau NL, Hayes JE. 2018. Self-reported smoking status, TAS2R38 variants, and propylthiouracil phenotype: an exploratory crowdsourced cohort study. Chem Senses. 43(3):617-625. 

    Di Pizio A, Ben Shoshan-Galeckki Y, Hayes JE, Niv MY. 2018. Bitter and sweet tasting molecules: it's complicated. Neurosci Lett. S0304-3940(18)30290-8.

    Moding, KJ, Ferrante MJ, Bellows LL, Bakke AJ, Hayes JE, Johnson SL. 2018. Variety and content of commercial infant and toddler vegetable products manufactured and sold in the United States. American Journal Clinical Nutrition. 107(4):576-583. 

    Bakke AJ, Stubbs CA, McDowell EH, Moding KJ, Johnson SJ, Hayes JE. 2018. Mary Poppins was right: adding small amounts of sugar or salt reduces the bitterness of vegetables. Appetite. 126: 90-101. 

    Fritts JR, Fort C, Quinn Corr A, Liang Q, Alla L, Cravener T, Hayes JE, Rolls BJ, D'Adamo C, Keller KL. 2018. Herbs and spices increase liking and preference for vegetables among rural high school students. Food Qual Pref. 68: 125-134. 

    Carney EM, Stein WM, Reigh NA, Gater FM, Bakke AJ, Hayes JE, Keller KL. 2018. Increasing flavor variety with herbs and spices improves relative vegetable intake in children who are propylthiouracil (PROP) tasters relative to nontasters. Physio Behav. 188: 48-57.


    Running CA, Hayes JE. 2017. Sip and Spit or Sip and Swallow: choice of method differentially alters taste intensity estimates across stimuli. Physiol Behav. 181:95-99.

    Elias RE, Hopfer H, Hofstaedter A, Hayes JE. Man vs. Machine: A Junior Level Laboratory Exercise Comparing Human and Instrumental Detection Limits. J Food Sci Edu. 16(3). 

    Reyes, MM, Castura J, Hayes JE. 2017. Characterizing Dynamic Sensory Properties of Nutritive and Nonnutritive Sweeteners with Temporal Check-All-That-Apply. J Sens Stud. 32(3).  

    Rolon ML, Bakke AJ, Coupland JN, Hayes JE, Roberts RF. 2017. Effect of fat content on physical properties and consumer acceptability of vanilla ice cream. J Dairy Sci. 100(7).  

    Johnson SL, Hayes JE. 2017. Developmental Readiness, Caregiver and Child Feeding Behaviors, and Sensory Science as a Framework for Feeding Young Children. Nutrition Today. 52(S): S30-40.  

    Hayes JE, Johnson SL. 2017. Sensory Aspects of Bitter and Sweet Tastes during Early Childhood. Nutrition Today. 52(S): S41-51.  

    Pickering GJ, Hayes JE. 2017. Influence of Biological, Experiential and Psychological Factors in Wine Preference Segmentation. Australian Journal of Grape and Wine Research. 23(2).  

    Tenney K, Hayes JE, Euston SR, Elias RJ, Coupland JN. 2017. Binding of Caffeine and Quinine by Whey Protein and the Effect on Bitterness. J Food Sci. 82: 509–516.  

    Bitzer ZT, Wopperer AL, Chrisfield BJ, Tao L, Cooper TK, Vanamala J, Elias RJ, Hayes JE. 2017. Soy protein concentrate mitigates markers of colonic inflammation and loss of gut barrier function in vitro and in vivo. J Nutrit Biochem. 40:201-208. 

    Nolden AA, Hayes JE. 2017. Perceptual and Affective Responses to Sampled Capsaicin Differ by Reported Intake. Food Qual Pref. 55:26-34. 


    Running CA, Ziegler GJ, Hayes JE. 2016. Degree of free fatty acid saturation influences chocolate rejection in human assessor. Chem Senses. Epub 20 Nov 2016. doi: 10.1093/chemse/bjw116

    Ziegler GR, Hayes JE, Lambert JD. Chocolate not necessarily healthier or tastier. PNAS. 113(42):e6318 

    Running CA, Hayes JE. Expectation and expectoration: information manipulation alters spitting volume, a common proxy for salivary flow. Physiol Behav. 167:180187. Epub 20 Sept 2016. doi: 10.1016/j.physbeh.2016.09.006

    Zaveri T, Running CA, Surapaneni L, Ziegler GR, Hayes JE. 2016. Innovative Sensory Methods to Assess Acceptability of Mixed Polymer Semisoft Ovules for Microbicide Applications. Drug Deliv Transl Res. 6(5), 551-564. Epub 29 June 2016. doi: 10.1007/s13346-016-0309-8

    Fleming EF, Ziegler GR, Hayes JE. Investigating Mixture Interactions of Astringent Stimuli using the Isobole Approach. Chem Senses. 41(7):601-610. Epub 1 June 2016. doi:10.1093/chemse/bjw064

    Perry DM, Hayes JE. 2016. Effects of Matrix Composition on Detection Threshold Estimates for methyl anthranilate and 2-aminoacetophenone. Foods. 5(2):35. Epub 17 May 2016. doi:10.3390/foods5020035

    Byrnes NK, Hayes JE. 2016. Behavioral measures of risk tasking, sensation seeking and sensitivity to reward may reflect different motivations for spicy food liking and consumption. Appetite. 103:411-422. Epub 29 April 2016. doi:10.1016/j.appet.2016.04.037.

    Fleming EF, Ziegler GR, Hayes JE. 2016. Salivary protein levels as a predictor of perceived astringency in model systems and solid foods. Physiol Behav. 163:56-63. Epub 27 April 2016. doi:10.1016/j.physbeh.2016.04.043

    Primrose RJ, Zaveri T, Bakke AJ, Ziegler GR, Moskowitz HR, Hayes JE. 2016. Drivers of vaginal drug delivery system acceptability from internet-based conjoint analysis. PLoS One. 11(3): e0150896. Epub 21 Mar 2016. doi 10.1371/journal.pone.0150896

    Sanyal S, O'Brien SM, Hayes JE, Feeney EL. 2016. TongueSim: Development of an automated method for rapid assessment of fungiform papillae density for taste research. Chem Senses. Epub 17 Feb 2016. doi: 10.1093/chemse/bjw008.

    Nolden AA, McGeary JE, Hayes JE. 2016. Differential bitterness in capsaicin, piperine, and ethanol associates with polymorphisms in multiple bitter taste receptor genes. Physiol Behav. 156:117-27. Epub 2016 Jan 16. doi: 10.1016/j.physbeh.2016.01.017

    Bakke AJ, †Shehan C, Hayes JE. 2016. Type of milk typically consumed, and stated preference, but not health consciousness affect revealed preferences for fat in milk. Food Qual Pref. 49:92-99. Epub 8 Dec 2015. doi:10.1016/j.foodqual.2015.12.001


    Nolden AA, Hayes JE. 2015. Perceptual qualities of ethanol depend on concentration, and variation in these percepts associates with drinking frequency. Chemosen Percept.  8(3):149-157. Epub 19 Sept 2015. doi:10.1007/s12078-015-9196-5

    Kelley K, Primrose RJ, Crassweller R, Hayes JE, Marini R. 2015. Consumer Peach Preferences and Purchasing Behavior: a mixed methods study. Journal of the Science of Food and Agriculture. In press. Epub 6 Aug 2015 doi:10.1002/jsfa.7365

    Byrnes NK, Loss C, Hayes JE. 2015. Perception of chemesthetic stimuli in groups who differ by food involvement and culinary experience. Food Qual Pref. 46: 142-150. Epub 27 July 2015. doi: 10.1016/j.foodqual.2015.07.017

    Webb J, Bolhuis D, Cicerale S, Hayes JE, Keast RSJ. 2015. The relationships between common measurements of taste function. Chemo Percept. 8(1):11-18. Epub 5 Jun 2015. doi: 10.1007/s12078-015-9183-x

    Fleming EF, Ziegler GR, Hayes JE. 2015. Check-All-That-Apply (CATA), Sorting, and Polarized Sensory Positioning (PSP) with Astringent Stimuli. Food Qual Prefer 45: 41-49. Epub 1 June 2015. doi:10.1016/j.foodqual.2015.05.004

    Hayes JE, Feeney EF, Nolden AA, McGeary JE. 2015. Quinine bitterness and grapefruit liking associate with allelic variants in TAS2R31. Chemical Senses. 40(6):437-443. Epub 29 May 2015. doi: 10.1093/chemse/bjv027.

    Byrnes NK, Nestrud M Hayes JE. 2015. Perceptual mapping of chemesthetic stimuli in naïve assessors. Chemo Percept. 8(1):19-32. Epub 22 March 2015. doi:10.1007/s12078-015-9178-7.

    Li B, Hayes JE, Ziegler GR. 2015. Maximizing overall liking results in a superior product to minimizing deviations from ideal ratings: an optimization case study with coffee-flavored milk. Food Qual Prefer. 42:27-36. Epub 20 Jan 2015.

    Byrnes NK, Hayes JE. 2015. Gender differences in the influence of personality traits on spicy food liking and intake. Food Qual Prefer. 42:12-19. Epub 9 Jan 2015. doi:10.1016/j.foodqual.2015.01.002

    Antenucci RG, Hayes JE. 2015. Non-nutritive sweeteners are not supernormal stimuli. Int J Obesity. 39:254-259. Epub 22 July 2014. doi: 10.1038/ijo.2014.109


    Coupland JN, Hayes JE. 2014. Physical Approaches to Masking Bitter Taste: Lessons from Food and Pharmaceuticals. Pharmaceutical Research. 31(11):2921-2939. Epub 10 Sept 2014. doi:10.1007/s11095-014-1480-6

    Allen AL, McGeary JE, Hayes JE. 2014. Polymorphisms in TRPV1 and TAS2Rs associate with sensations from sampled ethanol. Alcoholism Clinical and Experimental Research. 38(10):2550-2560. Epub 25 Sept 2014.

    Zaveri T, Primrose R, Surapaneni L, Ziegler GR, Hayes JE. 2014. Firmness Perception Influences Women's Preferences for Vaginal Suppositories. Pharmaceutics. 6(3), 512-529; doi:10.3390/ pharmaceutics6030512

    Mahan ED, Zaveri T, Ziegler GR, Hayes JE. 2014. Relationships between perceptual attributes and rheology in over-the-counter vaginal products: a potential tool for microbicide development. PLoS One. 9(9): e105614. doi:10.1371/journal.pone.0105614.

    Zaveri T, Hayes JE, Ziegler GR. 2014. Release of Tenofovir from Carrageenan-based Vaginal Suppositories. Pharmaceutics. 6(3):366-77. Epub 4 July 2014. doi:10.3390/pharmaceutics6030366.

    Antenucci RG, Hayes JE. 2014. Non-nutritive sweeteners are not supernormal stimuli. International Journal of Obesity (Lond). Epub 22 July 2014. doi: 10.1038/ijo.2014.109

    Garneau NL, Nuessle TM, Sloan MM, Santorico SA, Coughlin BC, Hayes JE. 2014. Crowdsourcing taste research: genetic and phenotypic predictors of bitter taste perception as a model. Front Integr Neurosci. 8:33. Epub 27 May 2014. doi:10.3389/fnint.2014.00033

    Li B, Hayes JE, Ziegler GR. 2014. Just-About-Right and ideal scaling provide similar insights into the influence of sensory attributes on liking. Food Qual Prefer. 37:71-78. Epub 10 May 2014. doi:/10.1016/j.foodqual.2014.04.01

    Feeney EL, Hayes JE. 2014. Regional differences in suprathreshold intensity for bitter and umami stimuli. Chemosensory Perception. Accepted 5 March 2014. Epub 13 May 2014. Doi:/10.1007/s12078-014-9166-3

    Li B, Hayes JE, Ziegler GR. 2014. Interpreting consumer preferences: physicohedonic and psychohedonic models yield different information in a coffee-flavored dairy beverage. Food Qual Prefer. 36(1):27-32. Epub 15 Mar 2014. doi:/10.1016/j.foodqual.2014.03.001

    Feeney EL, Hayes JE. 2014. Exploring associations between taste perception, oral anatomy and polymorphisms in the carbonic anhydrase (gustin) gene CA6. Physiol Behav. 128:148-154 Epub 15 Feb 2014. doi:10.1016/j.physbeh.2014.02.013

    Hayes JE, Raines CR, DePasquale DA, Cutter CN. 2014. Consumer acceptability of high hydrostatic pressure (HHP)-treated ground beef patties. LWT-Food Science and Technology. 56(1):207-210. Epub 21 Nov 2013. doi:10.1016/j.lwt.2013.11.014


    Rawal S, Hayes JE, Wallace MR, Bartoshuk LM, Duffy VB. 2013. Do polymorphisms in the TAS1R1 gene contribute to broader differences in human taste intensity? Chem Senses. 38(8):719-728. Epub 2 Sept 2013. doi:10.1093/chemse/bjt040

    Hayes JE, Feeney EL, Allen AL. 2013. Do polymorphisms in chemosensory genes matter for human ingestive behavior?. Food Qual Pref. 30(2): 202–216. doi:10.1016/j.foodqual.2013.05.013

    Harwood ML. Ziegler GR, and Hayes JE. 2013. Tolerance For High Flavanol Cocoa Powder in Semisweet Chocolate. Nutrients. 5(6): 2258-2267. Epub 21 June 2013.

    Allen A, McGeary JE, Hayes JE. 2013. Rebaudioside A and Rebaudioside D bitterness do not covary with Acesulfame K bitterness or polymorphisms in TAS2R9 and TAS2R31. Chemos Percept. 6(3):109-117. doi: 10.1007/s12078-013-9149-9.

    Harwood ML. Loquasto J, Roberts RF, Ziegler GR, and Hayes JE. 2013. Bitterness tolerance in chocolate ice cream. J Dairy Sci. Epub 13 June 2013. doi:10.3168/jds.2013-6715.

    Allen A, McGeary JE, Knopik V, Hayes JE. Bitterness of the non-nutritive sweetener Acesulfame Potassium varies with polymorphisms in TAS2R9 and TAS2R31. Chemical Senses. 38(5):379-389. Epub 18 Apr 2013

    Li B, Zaveri T, Ziegler GR, and Hayes JE. 2013. Shape of vaginal suppositories affects willingness-to-try and preference. Antiviral Research. Epub 29 Dec 2012. doi:10.1016/j.antiviral.2012.12.024

    Li B, Zaveri T, Ziegler GR, and Hayes JE. 2013. User preferences in a carrageenan-based vaginal drug delivery system. PLoS One. 8(1). Epub 24 Jan 2013. doi: 10.1371/journal.pone.0054975

    Sharafi M, Hayes JE, and Duffy VB. 2013. Masking vegetable bitterness to improve palatability depends on vegetable type and tastephenotype. Chemosensory Perception. Epub 28 Dec 2012. doi:10.1007/s12078-012-9137-5

    Byrnes NK, and Hayes JE. 2013. Personality factors predict spicy food liking and intake. Food Qual Pref. 28(1): 213-221. Epub 4 Oct 2012.doi:10.1016/j.foodqual.2012.09.008

    Hayes JE, Allen AL, and Bennett SM. 2013. Direct comparison of the generalized Visual Analog Scale (gVAS) and general Labeled Magnitude Scale (gLMS). Food Qual Pref. 28(1): 36-44. Epub 10 Aug 2012.doi:10.1016/j.foodqual.2012.07.012 


    Bennett SM, Zhou L, and Hayes JE. 2012. Using milk fat to reduce the irritation and bitter taste of ibuprofen. Chemosensory Perception.5(3-4):231-236. Epub 1 May 2012. doi: 10.1007/s12078-012-9128

    Harwood ML. Ziegler GR, and Hayes JE. 2012. Rejection Thresholds in Solid Chocolate-Flavored Compound Coating. J Food Sci. 77(10):S390-S393. Epub 27 Aug 2012. doi: 10.1111/j.1750-3841.2012.02889.x

    McGeary JE, Knopik VS, Hayes JE, Palmer RH, Monti P, Kalman D. Predictors of relapse in a bupropion trial for smoking cessation in recently-abstinent alcoholics: Preliminary results using an aggregate genetic risk score. Substance Abuse Research and Treatment. In press

    Bennett SM, Zhou L, and Hayes JE. Using milk fat to reduce the irritation and bitter taste of ibuprofen. Chemosensory Perception. Epub 1 May 2012. doi: 10.1007/s12078-012-9128-6.

    Harwood ML. Ziegler GR, and Hayes JE. 2012. Rejection Thresholds in Chocolate Milk: Evidence for Segmentation. Food Qual Pref. 26(1): 128­133. Epub 21 Apr 2012, doi: 10.1016/j.foodqual.2012.04.009.

    Bennett SM, Hayes JE. 2012. Differences in the chemesthetic sub-qualities of capsaicin, ibuprofen and olive oil. Chemical Senses. 37(5): 471-478 Epub 25 Jan 2012 doi:10.1093/chemse/bjr129.

    Harrington HL, Kennedy KE, Sharafi M, Hayes JE, Duffy VB. 2012. Otitis media exposure associates with dietary preference and adiposity: A community-based observational study of at-risk preschoolers. Physiol Behav. 106(2):264-271. Epub 6 Feb 2012, doi: 10.1016/j.physbeh.2012.01.021.

    Hayes JE, Pickering GJ. 2012. Wine expertise predicts taste phenotype. Am J Enol Viti. 63(1):80-84. Epub 2011 October, doi: 10.5344/ajev.2011.11050.


    Hayes JE, Keast RSJ. 2011. Two decades of supertasting: where do we stand?
    Physiol Behav. 104(5): 1072-1074. Epub 2011 Aug 7,

    Hayes JE, Keast RSJ. Two decades of supertasting: where do we stand? Physiol Behav. In press. Epub 2011 Aug 7, doi:10.1016/j.physbeh.2011.08.003

    Hayes JE, DePasquale DA, Moser SE. 2011. Asymmetric dominance as a potential source of bias in hedonic testing. Food Quality and Preference. 22(6):559-566. Epub 2011 Mar 21. doi:10.1016/j.foodqual.2011.03.006

    Mahan ED, Morrow KN, Hayes JE. 2011. Quantitative perceptual differences among over-the-counter vaginal products using a standardized methodology: implications for microbicide development. Contraception. 84(2):184-93. Epub 2011 Jan 10, doi:10.1016/j.contraception.2010.11.012

    Hayes JE, Knopik VS, et al. 2011. Allelic variation in TAS2R bitter receptor genes associates with variation in sensations from and ingestive behaviors towards common bitter beverages in adults. Chem Senses. 36(3):311-9. Epub 2010 Dec 16. doi:10.1093/chemse/bjq132


    Duffy VB, Hayes JE, et al. 2010. Vegetable intake in college-aged adults is explained by oral sensory phenotypes and TAS2R38 genotype. Chemosensory Perception. 3(3):137-148. Epub 2010 Aug 1. doi:10.1007/s12078-010-9079-8

    Duffy VB, Hayes JE, et al. 2010. ³Vegetable intake in college-aged adults is explained by oral sensory phenotypes and TAS2R38 genotype." Chemosensory Perception. Epub 2010 Aug 1. doi:10.1007/s12078-010-9079-8

    Hayes JE, Sullivan BS, Duffy VB. 2010. "Explaining variability in sodium intake through oral sensory phenotype, salt sensation and liking." Physiol Behav. 100(4): 369-380. Epub 2010 April 7,  doi:10.1016/j.physbeh.2010.03.017.

    Hayes JE. 2010. "Response to Lack of Relation Between Bitter Taste Receptor TAS2R38 and BMI in Adults." Obesity. 18, 433-433 (23 February 2010) doi:10.1038/oby.2009.351.

    2009 and earlier

    Duffy VB, Hayes JE, Sullivan BS, Faghri P. 2009. "Surveying Food/Beverage Liking: A Tool for Epidemiological Studies to Connect Chemosensation with Health Outcomes." Ann NY Acad Sci. 1170:558-568. doi: 10.1111/j.1749-6632.2009.04593.x.

    Hayes, JE and Duffy VB. 2008. "Oral sensory phenotype identifies level of sugar and fat required for maximal liking." Physiology & Behavior 95(1-2): 77-87. Epub 2008 May 2, doi:10.1016/j.physbeh.2008.04.023.

    Hayes JE, Bartoshuk LM, Kidd JK, Duffy VB. 2008. "Supertasting and PROP bitterness depends on more than the TAS2R38 gene." Chemical Senses 33(3): 255-265. Epub 2008 Jan 21, doi:10.1093/chemse/bjm084.

    Hayes, JE. 2008. "Transdisciplinary perspectives on Sweetness." Chemosensory Perception 1(1):48-57. Epub 2007 December 11, doi: 10.1007/s12078-007-9003-z.

    Hayes JE, Duffy VB. 2007. "Revisiting sugar-fat mixtures: sweetness and creaminess vary with phenotypic markers of oral sensation." Chemical Senses 32(3): 225-236. Epub 2007 Jan 4, doi:10.1093/chemse/bjl050.

    Bartoshuk LM, Duffy VB, Hayes JE, Snyder DJ. 2006. "Psychophysics of sweet and fat perception in obesity: problems, solutions and new perspectives." Philos Trans R Soc Lond B Biol Sci 361(1471): 1137-48.

    Dinehart ME, Hayes JE, Bartoshuk LM, Lanier SL. & Duffy VB. 2005. "Bitter taste markers explain variability in vegetable sweetness, bitterness, and intake." Physiology & Behavior, 87(2): 304-13.

    Lanier SA, Hayes JE, Duffy VB. 2005. "Sweet and bitter tastes of alcoholic beverages mediate alcohol intake in of-age undergraduates." Physiology & Behavior 83(5): 821-831.

    Green BG, Hayes JE. 2004. "Individual Differences in Perception of Bitterness from Capsaicin, Piperine and Zingerone." Chemical Senses 29(1): 53-60.

    Green BG, Hayes JE. 2003. "Capsaicin as a probe of the relationship between bitter taste and chemesthesis." Physiology & Behavior 79(4-5): 811-821.

    Horne J, Hayes J, Lawless HT. 2002. "Turbidity as a measure of salivary protein reactions with astringent substances." Chemical Senses 27(7): 653-9.

    Invited Book Chapters

    Duffy VB, Hayes JE, Bartoshuk LM, Snyder DJ. “Taste: vertebrates – Psychophysics." In: Reference Module in Neuroscience and Biobehavioral Psychology. 2017. ISBN 9780128093245.

    Harwood ML, Hayes JE. “Sensory Evaluation of chocolate and cocoa products" In: Beckett's Industrial Chocolate Manufacture and Use. 2017. ST Beckket, MS Fowler, GR Ziegler (eds). John Wiley & Sons..

    Running CA, Hayes JE. “Individual Differences in Multisensory Flavor Perception" In: Multisensory Flavor Perception. 2016. B. Piqueras-Fiszman and C. Spence. (eds). Elsevier.

    Hayes JE. “Pungency and burn: historical perspectives, word usage, and temporal characteristics" In: Chemesthesis: chemical touch in food and eating. 2016. S. McDonald, D. Bolliet, J.E. Hayes (eds). Wiley.

    Hayes JE. “Measuring sensory perception in relation to actual consumer behaviour" In: Rapid Sensory Profiling Techniques. 2015. J. Delarue, J. B. Lawlor and M. Rogeaux (eds). Woodhead Publishing. Cambridge. 53-69. ISBN 978-1782422488

    Prescott J, Hayes JE, Byrnes NK. “Sensory science" In: Encyclopedia of Agriculture and Food Systems. 2014. NK Van Alfen (editor-in-chief). Elsevier. San Diego. 80-101. ISBN 978-0-444-52512-3

    Corwin RL, Hayes JE, “Are sugars addictive? Perspectives for practitioners" In: Fructose, High Fructose Corn Syrup, Sucrose, and Health. James M. Rippe (editor). Springer. New York. 199-215. ISBN 978-1-4899-8077-9

    Duffy VB, Hayes JE, “Smell, taste, and oral somatosensation: age-related changes and nutritional implications" In: Geriatric Nutrition: the Health Professional's Handbook. 2013. 4th ed. Jones & Bartlett Learning. Burlington MA. ISBN 9780763782627.

    Duffy VB, Hayes JE, Bartoshuk LM, Snyder DJ. “Taste: vertebrate psychophysics." In: Encyclopedia of Neuroscience. 2009. Squire L (ed in chief). Academic Press. Oxford. 881-885. doi:10.1016/B978-008045046-9.01674-0.

    Snyder DJ, Duffy VB, Hayes JE, Bartoshuk LM. Propylthiouracil (PROP) taste. In: The Senses: A Comprehensive Reference. 2008. Basbaum et al (eds). Academic Press. New York. Vol 4, p. 391-399. doi: 10.1016/B978-012370880-9.00093-1.

    Duffy VB, Hayes JE, Dinehart ME. “Genetic differences in sweet taste perception." In: Optimising sweet taste in foods. 2006. Spillane WJ (ed). Woodhead Publishing Ltd. Cambridge.