Flavor: The Science of Our Most Neglected Sense - Bob Holmes 2017


Page numbers listed correspond to the print edition of this book. You can use your device’s search function to locate particular terms in the text.


4 Richard Wrangham argues: Richard Wrangham, Catching Fire: How Cooking Made Us Human (New York: Basic Books, 2009): 105—127.

4 spices have antibacterial properties: Paul W. Sherman and Jennifer Billing, “Darwinian Gastronomy: Why We Use Spices,” BioScience 49 (1999): 453—463.

5 more brain systems: Gordon M. Shepherd, “Neuroenology: How the Brain Creates the Taste of Wine,” Flavour 4 (2015): 19, doi:10.1186/s13411-014-0030-9.

6 Vegemite: Paul Rozin and Michael Siegal, “Vegemite as a Marker of National Identity,” Gastronomica 3, no. 4 (2003): 63—67.

6 holidays, sex, and family time: J. Westenhoefer and V. Pudel, “Pleasure from Food: Importance for Food Choice and Consequences of Deliberate Restriction,” Appetite 20 (1993): 246.

8 English speakers generally use taste: Paul Rozin, “’Taste-Smell Confusions’ and the Duality of the Olfactory Sense,” Perception and Psychophysics 31 (1982): 397—401.

11 Bush told reporters: Maureen Dowd, “’I’m President,’ So No More Broccoli!,” New York Times, March 23, 1990,


14 Bartoshuk who first suggested: Linda M. Bartoshuk, Valerie B. Duffy, and Inglis J. Miller, “PTC/PROP Tasting: Anatomy, Psychophysics, and Sex Effects,” Physiology & Behavior 56 (1994): 1165—1171.

17 lose those extraneous tastes: Alexander A. Bachmanov et al., “Genetics of Taste Receptors,” Current Pharmaceutical Design 20 (2014): 2669—2683.

17 vampire bats: Wei Hong and Huabin Zhao, “Vampire Bats Exhibit Evolutionary Reduction of Bitter Taste Receptor Genes Common to Other Bats,” Proceedings of the Royal Society B (2014), doi:10.1098/rspb.2014.1079.

23 104 diverse bitter-tasting chemicals: Wolfgang Meyerhof et al., “The Molecular Receptive Ranges of Human TAS2R Bitter Taste Receptors,” Chemical Senses 35 (2010): 157—170.

26 published a letter: Robert Ho Man Kwok, “Chinese-Restaurant Syndrome,” New England Journal of Medicine 278 (1968): 796.

26 picked up the story: For a good treatment of the history of Chinese restaurant syndrome, see Ian Mosby, “’That Won-Ton Soup Headache’: The Chinese Restaurant Syndrome, MSG and the Making of American Food, 1968—1980,” Social History of Medicine (2009): 133—151, doi:10.1093/shm/hkn098.

27 fifty-eight million pounds: Ibid., 7.

27 most damning evidence: L. Tarasoff and M. F. Kelly, “Monosodium L-Glutamate: A Double-Blind Study and Review,” Food and Chemical Toxicology 31 (1993): 1019—1035.

27 when researchers looked back: Ibid.

29 discovered by accident: Anonymous, “The Inventor of Saccharine,” Scientific American, July 17, 1886, 36.

29 Cyclamate: Deborah Jean Warner, Sweet Stuff: An American History of Sweeteners from Sugar to Sucralose (Lanham, MD: Rowman & Littlefield, 2011), 195, accessed via Google Books, March 29, 2016.

29 Aspartame: Robert H. Mazur, “Discovery of Aspartame,” in Lewis D. Stegink and L. J. Filer, Jr., eds., Aspartame: Physiology and Biochemistry (New York: Marcel Dekker, 1984), 4.

29 Sucralose: Burkhard Bilger, “The Search for Sweet,” The New Yorker, May 22, 2006, 40.

30 Pepsi is about 11 percent: Daniel Engber, “The Quest for a Natural Sugar Substitute,” New York Times Magazine, January 1, 2014,

30 its own distinctive timing: Paul A. S. Breslin and Alan C. Spector, “Mammalian Taste Perception,” Current Biology 18 (2008): R153.

30 ten seconds later: Engber, “Quest for a Natural Sugar.”

30 four seconds longer: Ibid.

31 9 grams of salt daily: S. L. Drake and M. A. Drake, “Comparison of Salty Taste and Time Intensity of Sea and Land Salts from around the World,” Journal of Sensory Studies 26 (2010): 25.

31 from processed foods: Marjorie Ellin Doyle and Kathleen A. Glass, “Sodium Reduction and Its Effect on Food Safety, Food Quality, and Human Health,” Comprehensive Reviews in Food Science and Food Safety 9 (2010): 44—56.

31 high blood pressure: Ibid., 45.

32 they taste saltier: Tassyana Vieira Marques Freire et al., “Salting Potency and Time-Intensity Profile of Microparticulated Sodium Chloride in Shoestring Potatoes,” Journal of Sensory Studies 30 (2015): 1—9.

33 bitter-tasting medicines: Adam A. Clark, Stephen B. Liggett, and Steven D. Munger, “Extraoral Bitter Taste Receptors as Mediators of Off-Target Drug Effects,” FASEB Journal 26 (2012): 4827—4831.

34 more sinus infections: Robert J. Lee and Noam A. Cohen, “The Emerging Role of the Bitter Taste Receptor T2R38 in Upper Respiratory Infection and Chronic Rhinosinusitis,” American Journal of Rhinology and Allergy 27 (2013): 283—286.

35 receptors on our taste buds: Robin M. Tucker, Richard D. Mattes, and Cordelia A. Running, “Mechanisms and Effects of ’Fat Taste’ in Humans,” BioFactors 40 (2014): 313—326.

35 a distinct taste: Cordelia A. Running, Bruce A. Craig, and Richard D. Mattes, “Oleogustus: The Unique Taste of Fat,” Chemical Senses 40 (2015), 507—516.

35 oleogustus: Ibid.

36 a taste for calcium: Michael G. Tordoff et al., “T1R3: A Human Calcium Taste Receptor,” Scientific Reports 2 (2012): 496, doi:10.1038/srep00496.

36 for carbon dioxide: Jayaram Chandrashekar et al., “The Taste of Carbonation,” Science 326 (2009): 443—445.

36 a taste for starch: Breslin and Spector, “Mammalian Taste Perception,” R149.

37 calcium-sensing receptor: Motonaka Kuroda and Naohiro Miyamura, “Mechanism of the Perception of ’Kokumi’ Substances and the Sensory Characteristics of the ’Kokumi’ Peptide, Gamma-Glu-Val-Gly,” Flavour 4 (2015): 11, doi:10.1186/2044-7248-4-11.

37 interact with one another: Russell S. J. Keast and Paul A. S. Breslin, “An Overview of Binary Taste-Taste Interactions,” Food Quality and Preference 14 (2002): 117.

37 ability to taste PROP: Bernd Bufe et al., “The Molecular Basis of Individual Differences in Phenylthiocarbamide and Propylthiouracil Bitterness Perception,” Current Biology 15 (2005): 322—327.

38 That’s probably why: Bartoshuk, Duffy, and Miller, “PTC/PROP Tasting.”

39 support that hunch: For example, John E. Hayes and Valerie B. Duffy, “Revisiting Sugar-Fat Mixtures: Sweetness and Creaminess Vary with Phenotypic Markers of Oral Sensation,” Chemical Senses 32 (2007): 225—236.

39 fail to find a link: For example, Mary E. Fischer et al., “Factors Related to Fungiform Papillae Density: The Beaver Dam Offspring Study,” Chemical Senses 38 (2013): 669—677; Nicole L. Garneau et al., “Crowdsourcing Taste Research: Genetic and Phenotypic Predictors of Bitter Taste Perception as a Model,” Frontiers in Integrative Neuroscience 8 (2014): 33, doi:10.3389/fnint.2014.00033.

39 gustin might be involved: Melania Melis et al., “The Gustin (CA6) Gene Polymorphism, rs2274333 (A/G) as a Mechanistic Link between PROP Tasting and Fungiform Taste Papilla Density and Maintenance,” PLoS One 8 (2013): e74151, doi:10.1371/journal.pone.0074151.

40 affects sweet perception: Alexey A. Fushan et al., “Allelic Polymorphism within the TAS1R3 Promoter Is Associated with Human Taste Sensitivity to Sucrose,” Current Biology 19 (2009): 1288—1293.

42 two kinds of supertasters: Natalia V. Ullrich et al., “PROP Taster Status and Self-Perceived Food Adventurousness Influence Food Preferences,” Journal of the American Dietetic Association 104 (2004): 543—549.


48 unique pattern of molecular vibrations: For the most detailed exposition of this point of view, see Luca Turin, The Secret of Scent: Adventures in Perfume and the Science of Smell (London: Faber and Faber, 2006).

49 a different chord: This apt analogy is not mine, alas. I first encountered it on the Food Sommelier website,

49 used it in their key paper: Linda Buck and Richard Axel, “A Novel Multigene Family May Encode Odorant Receptors: A Molecular Basis for Odor Recognition,” Cell 65 (1991): 183.

50 But a closer look: This paragraph follows Avery Gilbert, What the Nose Knows: The Science of Scent in Everyday Life (New York: Crown, 2008): 2—4.

53 413 odor receptors: Tsviya Olender et al., “Personal Receptor Repertoires: Olfaction as a Model,” BMC Genomics 13 (2012): 414, doi:10.1186/1471-2164-13-414.

55 ORs all over the place: Ester Feldmesser et al., “Widespread Ectopic Expression of Olfactory Receptor Genes,” BMC Genomics 7 (2006): 121, doi:10.1186/1471-2164-7-121.

55 combine ethyl isobutyrate: E. Le Berre et al., “Just Noticeable Differences in Component Concentrations Modify the Odor Quality of a Blending Mixture,” Chemical Senses 33 (2008), 389—395.

55 one part geraniumy: C. Masanetz, H. Guth, and W. Grosch, “Fishy and Hay-like Off-flavours of Dry Spinach,” Zeitschrift für Lebensmitteluntersuchung und -Forschung A 206 (1998): 108—113.

56 a trillion different odor objects: C. Bushdid et al., “Humans Can Discriminate More Than 1 Trillion Olfactory Stimuli,” Science 243 (2014): 1370—1372.

56 treated with caution: Richard C. Gerkin and Jason B. Castro, “The Number of Olfactory Stimuli That Humans Can Discriminate Is Still Unknown,” eLife 4 (2015): e08127, doi:10.7554/eLife.08127.

57 a terrific book: Gordon M. Shepherd, Neurogastronomy (New York: Columbia University Press, 2012).

57 asked her to close her eyes: Yaara Yeshurun and Noam Sobel, “An Odor Is Not Worth a Thousand Words: From Multidimensional Odors to Unidimensional Odor Objects,” Annual Review of Psychology 61 (2010): 226.

58 straight to a neurologist: The colleague is Jay Gottfried of Northwestern University. See Greg Miller, “What’s Up with That: Why Are Smells So Difficult to Describe in Words?” Wired, November 11, 2014,

59Big Red gum”: This quote is from Asifa Majid and Niclas Burenhult, “Odors Are Expressible in Language, as Long as You Speak the Right Language,” Cognition 130 (2014): 266—270.

59 quick and consistent: Ibid.

60 wine experts’ noses are no better: Wendy V. Parr, David Heatherbell, and K. Geoffrey White, “Demystifying Wine Expertise: Olfactory Threshold, Perceptual Skill and Semantic Memory in Expert and Novice Wine Judges,” Chemical Senses 27 (2002): 747—755.

60 fell off dramatically: D. G. Laing and G. W. Francis, “The Capacity of Humans to Identify Odors in Mixtures,” Physiology & Behavior 46 (1989): 809—814.

60 Later studies have confirmed: Anthony Jinks and David G. Laing, “A Limit in the Processing of Components in Odour Mixtures,” Perception 28 (1999): 395—404.

61 ambrosial and stench: Stanley Finger, Origins of Neuroscience: A History of Explorations into Brain Function (Oxford, UK: Oxford University Press, 2001), 178.

61 The Suya: Constance Classen, David Howes, and Anthony Synnott, Aroma: The Cultural History of Smell (London: Routledge, 1994), 100—101.

62 The Serer-Ndut: Ibid., 102—104.

64 chocolate-tracking experiment: Jess Porter et al., “Mechanisms of Scent-Tracking in Humans,” Nature Neuroscience 10 (2007): 27—29.

66 don’t get any better: Lee Sela and Noam Sobel, “Human Olfaction: A Constant State of Change-Blindness,” Experimental Brain Research 205 (2010): 13—29.

66 smell your hand: Idan Frumin et al., “A Social Chemosignaling Function for Human Handshaking,” eLife 4 (2015): e05154, doi:10.7554/eLife.05154.

66 Sobel told a reporter: Catherine de Lange, “After Handshakes, We Sniff People’s Scent on Our Hand,” New Scientist, March 3, 2015,

67 one famous experiment: Daniel J. Simons and Daniel T. Levin, “Failure to Detect Changes to People During a Real-World Interaction,” Psychonomic Bulletin and Review 5 (1998): 644—649.

67 change blindness: Sela and Sobel, “Human Olfaction.”

68 forms an air curtain: Rui Ni et al., “Optimal Directional Volatile Transport in Retronasal Olfaction,” Proceedings of the National Academy of Sciences 112 (2015): 14700—14704.

68 according to Shepherd: Shepherd, Neurogastronomy, 19—27.

70 thresholds tend to be lower: Viola Bojanowski and Thomas Hummel, “Retronasal Perception of Odors,” Physiology & Behavior 107 (2012): 484—487.

71 smells different to each nostril: Noam Sobel et al., “The World Smells Different to Each Nostril,” Nature 402 (1999): 35.

72 30 percent of our odor receptors: Joel D. Mainland et al., “The Missense of Smell: Functional Variability in the Human Odorant Receptor Repertoire,” Nature Neuroscience 17 (2014): 114—120.

74 genes help determine: Charles J. Wysocki and Gary K. Beauchamp, “Ability to Smell Androstenone Is Genetically Determined,” Proceedings of the National Academy of Sciences 81 (1984), 4899—4902.

76 can vary many thousandfold: Andreas Keller et al., “An Olfactory Demography of a Diverse Metropolitan Population,” BMC Neuroscience 13 (2012): 122, doi:10.1186/1471-2202-13-122.

76 nothing special: For example, Parr, Heatherbell, and White, “Demystifying Wine Expertise.”

77 easier to detect: David E. Hornung et al., “Effect of Nasal Dilators on Nasal Structures, Sniffing Strategies, and Olfactory Ability,” Rhinology 39 (2001): 84—87.

77 More than a thousand other genes: Ifat Keydar et al., “General Olfactory Sensitivity Database (GOSdb): Candidate Genes and Their Genomic Variations,” Human Mutation 34 (2012): 32—41.

77transforms my chamber pot”: Quoted in Marcia Levin Pelchat et al., “Excretion and Perception of a Characteristic Odor in Urine after Asparagus Ingestion: A Psychophysical and Genetic Study,” Chemical Senses 36 (2011): 9—17.

77 a pound of canned asparagus: M. Lison, S. H. Blondheim, and R. N. Melmed, “A Polymorphism of the Ability to Smell Urinary Metabolites of Asparagus,” British Medical Journal 281 (1980): 20—27.

78 OR2M7: Nicholas Eriksson et al., “Web-Based, Participant-Driven Studies Yield Novel Genetic Associations for Common Traits,” PLoS Genetics 6 (2010): e1000993, doi:10.1371/journal.pgen.1000993.

78 who really do produce odorless urine: Marcia Pelchat et al., “Excretion and Perception of a Characteristic Odor in Urine after Asparagus Ingestion: A Psychophysical and Genetic Study,” Chemical Senses 36 (2010): 9—17.

78 OR6A2 gene: Nicholas Eriksson et al., “A Genetic Variant near Olfactory Receptor Genes Influences Cilantro Preference,” Flavour 1 (2012): 22, doi:10.1186/2044-7248-1-22.

81 Smell-O-Vision: See Gilbert, What the Nose Knows: 155—163.

83 recent German study: Andreas Dunkel et al., “Nature’s Chemical Signatures in Human Olfaction: A Foodborne Perspective for Future Biotechnology,” Angewandte Reviews 53 (2014): 7124—7143.


87 the receptor for capsaicin: Michael J. Caterina et al., “The Capsaicin Receptor: A Heat-Activated Ion Channel in the Pain Pathway,” Nature 389 (1997): 816—824.

88 extra-virgin olive oil: Catherine Peyrot des Gachons et al., “Unusual Pungency from Extra-Virgin Olive Oil Is Attributable to Restricted Spatial Expression of the Receptor of Oleocanthal,” Journal of Neuroscience 31 (2011): 999—1009.

95 report less burn: Pamela Dalton and Nadia Byrnes, “The Psychology of Chemesthesis: Why Would Anyone Want to Be in Pain?,” in Shane T. McDonald, David Bolliet, and John Hayes, eds., Chemesthesis: Chemical Touch in Food and Eating (Chichester, UK: Wiley, 2016), 8—31.

96 58 percent of our liking: Outi Tornwall et al., “Why Do Some Like It Hot? Genetic and Environmental Contributions to the Pleasantness of Oral Pungency,” Physiology & Behavior 107 (2012): 381—389.

96benign masochism”: Paul Rozin and Deborah Schiller, “The Nature and Acquisition of a Preference for Chili Pepper by Humans,” Motivation and Emotion 4 (1980): 77—101.

98 more likely to be sensation seekers: Nadia K. Byrnes and John E. Hayes, “Personality Factors Predict Spicy Food Liking and Intake,” Food Quality and Preference 28 (2013): 213—221.

98 an interesting pattern emerged: Nadia K. Byrnes and John E. Hayes, “Gender Differences in the Influence of Personality Traits on Spicy Food Liking and Intake,” Food Quality and Preference 42 (2015): 12—19.

99 fifty-hertz vibration: Nobuhiro Hagura, Harry Barber, and Patrick Haggard, “Food Vibrations: Asian Spice Sets Lips Trembling,” Proceedings of the Royal Society B 280 (2013): 1680, doi:10.1098/rspb.2013.1680.

100 blocks the flow of potassium: Kristin A. Gerhold and Diana M. Bautista, “Molecular and Cellular Mechanisms of Trigeminal Chemosensation,” Annals of the New York Academy of Sciences 1170 (2009): 184—189.

100 high-altitude climber: Mark Graber and Stephen Kelleher, “Side Effects of Acetazolamide: The Champagne Blues,” American Journal of Medicine 84 (1988): 979—980.

101 eliminating the bubbles: Paul M. Wise et al., “The Influence of Bubbles on the Perception Carbonation Bite,” PLoS One 8 (2013): e71488, doi:10.1371/journal.pone.0071488.

103 astringency built up: Catherine Peyrot des Gachons et al., “Opponency of Astringent and Fat Sensations,” Current Biology 22 (2012): R829—R830.

103 reported the first hints: Nicole Schöbel et al., “Astringency Is a Trigeminal Sensation That Involves the Activation of G Protein-Coupled Signaling by Phenolic Compounds,” Chemical Senses 39 (2014): 471—487.


109 sugar tasted sweeter: Richard J. Stevenson, John Prescott, and Robert A. Boakes, “Confusing Tastes and Smells: How Odours Can Influence the Perception of Sweet and Sour Tastes,” Chemical Senses 24 (1999): 627—635.

109 host of similar studies: For an overview of these, see Malika Auvray and Charles Spence, “The Multisensory Perception of Flavor,” Consciousness and Cognition 17 (2008): 1016—1031.

111 Pringles potato chips: Massimiliano Zampini and Charles Spence, “The Role of Auditory Cues in Modulating the Perceived Crispness and Staleness of Potato Chips,” Journal of Sensory Studies 19 (2004): 347—363.

111 sounds of a coffee maker: Klemens Michael Knöferle, “Acoustic Influences on Consumer Behavior: Empirical Studies on the Effects of In-Store Music and Product Sound,” (PhD dissertation, University of St. Gallen, 2011), 36,$FILE/dis3964.pdf.

112 when accompanied by the sea sounds: Charles Spence, Maya U. Shankar, and Heston Blumenthal, “’Sound Bites’: Auditory Contributions to the Perception and Consumption of Food and Drink,” in Francesca Bacci and David Melcher, eds., Art and the Senses (Oxford, UK: Oxford University Press, 2011), 225—226.

113 words like kiki: Alberto Gallace, Erica Boschin, and Charles Spence, “On the Taste of ’Bouba’ and ’Kiki’: An Exploration of Word-Food Associations in Neurologically Normal Participants,” Cognitive Neuroscience 2 (2011): 34—46.

113 ice cream called “Frosh”: Eric Yorkston and Geeta Menon, “A Sound Idea: Phonetic Effects of Brand Names on Consumer Judgments,” Journal of Consumer Research 31 (2004): 43—51.

114 the crockery: This aspect is discussed extensively in Charles Spence and Betina Piqueras-Fiszman, The Perfect Meal: The Multisensory Science of Food and Dining (Chichester, UK: Wiley, 2014): 109—143.

115 sweetness, but not its saltiness: J. A. Maga, “Influence of Color on Taste Thresholds,” Chemical Senses 1 (1974): 115—119.

115 had tinted it red: Gil Morrot, Frédéric Brochet, and Denis Dubourdieu, “The Color of Odors,” Brain and Language 79 (2001): 309—320.

116 three different rooms: Carlos Velasco et al., “Assessing the Influence of the Multisensory Environment on the Whisky Drinking Experience,” Flavour 2 (2013): 23, doi:10.1186/2044-7248-2-23.

116 ice hockey games: Corinna Noel and Robin Dando, “The Effect of Emotional State on Taste Perception,” Appetite 95 (2015): 89—95.

119 squirm-inducing experiment: Dana M. Small et al., “Differential Neural Responses Evoked by Orthonasal versus Retronasal Odorant Perception in Humans,” Neuron 47 (2005): 593—605.

121 Shepherd puts it best: Gordon M. Shepherd, Neurogastronomy (New York: Columbia University Press, 2012), ix (emphasis in original).

123 labeled as “body odor”: Ivan E. de Araujo et al., “Cognitive Modulation of Olfactory Processing,” Neuron 46 (2005): 671—679.

125 results were shocking: Robert T. Hodgson, “An Examination of Judge Reliability at a Major U.S. Wine Competition,” Journal of Wine Economics 3 (2008): 105—113.

125 other major wine competitions: Robert T. Hodgson, “An Analysis of the Concordance Among 13 U.S. Wine Competitions,” Journal of Wine Economics 4 (2009): 1—9.

126 not humanly possible to judge wines objectively: I owe this idea to Anna Katharine Mansfield, a wine researcher at Cornell University.

126 prefer cheaper wines: Robin Goldstein et al., “Do More Expensive Wines Taste Better? Evidence from a Large Sample of Blind Tastings,” Journal of Wine Economics 3 (2008): 1—9.

127 wines of varying price: Hilke Plassmann et al., “Marketing Actions Can Modulate Neural Representations of Experienced Pleasantness,” Proceedings of the National Academy of Sciences 105 (2008): 1050—1054.

128 The frontal operculum: Janina Seubert et al., “Superadditive Opercular Activation to Food Flavor Is Mediated by Enhanced Temporal and Limbic Coupling,” Human Brain Mapping 36 (2015): 1662—1676.

129 in monkey brains: Edmund T. Rolls et al., “Sensory-Specific Satiety: Food-Specific Reduction in Responsiveness of Ventral Forebrain Neurons after Feeding in the Monkey,” Brain Research 368 (1986): 79—86.

129 gradually switch their responses: Edmund T. Rolls et al., “Orbitofrontal Cortex Neurons: Role in Olfactory and Visual Association Learning,” Journal of Neurophysiology 75 (1996): 1970—1981.

131 smell more accurately: Jahan B. Jadauji et al., “Modulation of Olfactory Perception by Visual Cortex Stimulation,” Journal of Neuroscience 32 (2012): 3095—3100.


134 Sclafani offered rats: Catalina Pérez, François Lucas, and Anthony Sclafani, “Increased Flavor Acceptance and Preference Conditioned by the Postingestive Actions of Glucose,” Physiology & Behavior 64 (1998): 483—492.

136 learned which flavor delivered: Ivan E. de Araujo et al., “Metabolic Regulation of Brain Response to Food Cues,” Current Biology 23 (2013): 878—883.

137 An old study from the 1950s: James Olds and Peter Milner, “Positive Reinforcement Produced by Electrical Stimulation of Septal Area and Other Regions of Rat Brain,” Journal of Comparative and Physiological Psychology 47 (1954): 419—427.

138 conscious and unconscious valuations: Deborah W. Tang, Lesley K. Fellows, and Alain Dagher, “Behavioral and Neural Valuation of Foods Is Driven by Implicit Knowledge of Caloric Content,” Psychological Science 25 (2014): 2168—2176.

141 a glass of carrot juice: Julie A. Mennella, Coren P. Jagnow, and Gary K. Beauchamp, “Prenatal and Postnatal Flavor Learning by Human Infants,” Pediatrics 107 (2001): E88,

142 as infants liked it better: R. Haller et al., “The Influence of Early Experience with Vanillin on Food Preference Later in Life,” Chemical Senses 24 (1999): 465—467.

142 more accepting of vegetable flavors: Julie A. Mennella, “Ontogeny of Taste Preferences: Basic Biology and Implications for Health,” American Journal of Clinical Nutrition 99 (2014): 704S—711S.

143you are turning green”: Carol Zane Jolles, Faith, Food, and Family in a Yupik Whaling Community (Seattle: University of Washington Press, 2002), 284; cited in Sveta Yamin-Pasternak et al., “The Rotten Renaissance in the Bering Strait: Loving, Loathing, and Washing the Smell of Foods with a (Re)acquired Taste,” Current Anthropology 55 (2014): 619—646.

144 wearing latex gloves: Yamin-Pasternak et al., “Rotten Renaissance.”

144 didn’t respond the same way: Paul M. Wise et al., “Reduced Dietary Intake of Simple Sugars Alters Perceived Sweet Taste Intensity but Not Perceived Pleasantness,” American Journal of Clinical Nutrition 103 (2016): 50—60.

146 the renowned French Laundry: Thomas Keller, The French Laundry Cookbook (New York: Artisan, 1999): 14.

147 tomato soup aroma: Mariëlle Ramaekers et al., “Aroma Exposure Time and Aroma Concentration in Relation to Satiation,” British Journal of Nutrition 111(2014): 554—562.

147 some other young Dutch men: Anne G. M. Wijlens et al., “Effects of Oral and Gastric Stimulation on Appetite and Energy Intake,” Obesity 20 (2012): 2226—2232.

148 big squirts separated: Dieuwerke P. Bolhuis et al., “Both Longer Oral Sensory Exposure to and Higher Intensity of Saltiness Decrease Ad Libitum Food Intake in Healthy Normal-Weight Men,” Journal of Nutrition 141 (2011): 2242—2248.

149 eating pasta with a small spoon: Ana M. Andrade et al., “Does Eating Slowly Influence Appetite and Energy Intake When Water Intake Is Controlled?” International Journal of Behavioral Nutrition and Physical Activity 9 (2012): 135, doi:10.1186/1479-5868-9-135.

149 playing with texture: K. McCrickerd and C. G. Forde, “Sensory Influences on Food Intake Control: Moving beyond Palatability,” Obesity Reviews 17 (2015): 18—29.

149 soup more filling: Mieke J. I. Martens and Margriet S. Westerterp-Plantenga, “Mode of Consumption Plays a Role in Alleviating Hunger and Thirst,” Obesity 20 (2012): 517—524.

150 highly flavored vanilla custard: René A. de Wijk et al., “Food Aroma Affects Bite Size,” Flavour 1 (2012): 3, doi:10.1186/2044-7248-1-3.

150 saltier tomato soup: Bolhuis et al., “Longer Oral Sensory Exposure.”

150 a dozen rat-friendly flavors: Michael Naim et al., “Energy Intake, Weight Gain, and Fat Deposition in Rats Fed Flavored, Nutritionally Controlled Diets in a Multichoice (’Cafeteria’) Design,” Journal of Nutrition 115 (1985): 1447—1458.

151 scarf the stuff down anyway: Israel Ramirez, “Influence of Experience on Response to Bitter Taste,” Physiology & Behavior 49 (1991): 387—391.

152 any taste receptor or odor receptor genes: Adam E. Locke et al., “Genetic Studies of Body Mass Index Yield New Insights for Obesity Biology,” Nature 518 (2015): 197—206.

156 tells you nothing useful: Chih-Hung Shu et al., “The Proportion of Self-Rated Olfactory Dysfunction Does Not Change across the Life Span,” American Journal of Rhinology & Allergy 23 (2009): 413—416.

156 impairments of their sense of smell: Claire Murphy et al., “Prevalence of Olfactory Impairment in Older Adults,” JAMA 288 (2002): 2307—2312.

156 scratch-and-sniff smell survey: Charles J. Wysocki and Avery N. Gilbert, “National Geographic Smell Survey: Effects of Age Are Heterogeneous,” Annals of the New York Academy of Sciences 561 (1989): 12—28.

158 responded to both: Nancy E. Rawson et al., “Age-Associated Loss of Selectivity in Human Olfactory Sensory Neurons,” Neurobiology of Aging 33 (2012): 1913—1919.

158 four times as likely to die: Jayant M. Pinto et al., “Olfactory Dysfunction Predicts 5-Year Mortality in Older Adults,” PLoS One 9 (2014): e107541, doi:10.1371/journal.pone.0107541.

159 depression and anxiety: Carl M. Philpott and Duncan Boak, “The Impact of Olfactory Disorder in the United Kingdom,” Chemical Senses 39 (2014): 711—718.

159 linked to other health problems: Nicole Toussaint et al., “Loss of Olfactory Function and Nutritional Status in Vital Older Adults and Geriatric Patients,” Chemical Senses 40 (2015): 197—203.

160 might improve with practice: Thomas Hummel et al., “Effects of Olfactory Training in Patients with Olfactory Loss,” Laryngoscope 119 (2009): 496—499.

161 Mark Friedman thinks: David S. Ludwig and Mark I. Friedman, “Increasing Adiposity: Cause or Consequence of Overeating?” Journal of the American Medical Association 311 (2014): 2167—2168.

162 Dana Small thinks: Martin G. Myers Jr. et al.,”Obesity and Leptin Resistance: Distinguishing Cause from Effect,” Trends in Endocrinology and Metabolism 21 (2010): 643—651.

162 containers of sugary or fatty food: Michael G. Tordoff, “Obesity by Choice: The Powerful Influence of Nutrient Availability on Nutrient Intake,” American Journal of Physiology: Regulatory, Integrative, and Comparative Physiology 282 (2002): R1536—R1539.


166 more than $10 billion worth of flavorings:

187 Virgin Mary in a grilled-cheese sandwich: Jessica Firger, “See the Virgin Mary on Toast? No, You’re Not Crazy,” CBS News, May 4, 2014, The phenomenon has been studied scientifically: Jiangang Liu et al., “Seeing Jesus in Toast: Neural and Behavioral Correlates of Face Pareidolia,” Cortex 53 (2014): 60—77.

191 many other languages have fewer: Joseph Henrich, The Secret of Our Success: How Culture Is Driving Human Evolution, Domesticating Our Species, and Making Us Smarter (Princeton, NJ: Princeton University Press, 2016), 240.

194 Australian chemistry teacher: See, for example, James Kennedy, “Ingredients of an All-Natural Banana,”

196the Dorito effect”: Mark Schatzker, The Dorito Effect: The Surprising New Truth about Food and Flavor (New York: Simon & Schuster, 2015).

198note-by-note cooking”: Hervé This, Note-by-Note Cooking: The Future of Food (New York: Columbia University Press, 2014).

198 BBC news report: “Is This What We’ll Eat in the Future?”, video of Hervé This, BBC News, November 6, 2013,

198 told one reporter: Wendell Steavenson, “Hervé This: The World’s Weirdest Chef,” Prospect, September 2014,

199dirac”: Hervé This, “Three Recipes for Note by Note Cooking,” La Cuisine Note à Note (blog), November 20, 2014,


205 40 percent lower than they used to be: Donald R. Davis, “Declining Fruit and Vegetable Nutrient Composition: What Is the Evidence?” Hort-Science 44 (2009): 15—19.

206 chose sixty-six varieties: Denise Tieman et al., “The Chemical Interactions Underlying Tomato Flavor Preferences,” Current Biology 22 (2012): 1035—1039.

206 twice as sweet: Linda M. Bartoshuk and Harry J. Klee, “Better Fruits and Vegetables through Sensory Analysis,” Current Biology 23 (2013): R374—R378.

208 to essential human nutrients: Stephen A. Goff and Harry J. Klee, “Plant Volatile Compounds: Sensory Cues for Health and Nutritional Value?” Science 311 (2006): 815—819.

210 tested what made for a tasty tomato: Tieman et al., “Tomato Flavor Preferences.”

213 looked at the volatiles in strawberries: Michael L. Schwieterman et al., “Strawberry Flavor: Diverse Chemical Compositions, a Seasonal Influence, and Effects on Sensory Perception,” PLoS One 9 (2014): e88446, doi:10.1371/journal.pone.0088446.

213 a single gene variant: Alan H. Chambers et al., “Identification of a Strawberry Flavor Gene Candidate Using an Integrated Genetic-Genomic-Analytical Chemistry Approach,” BMC Genomics 15 (2014): 217, doi:10.1186/1471-2164-15-217.

217 description that emerges: Wendy V. Parr et al., “Perceived Minerality in Sauvignon Wines: Influence of Culture and Perception Mode,” Food Quality and Preference 41 (2015): 121—132.

218 ten times as much thiol: W. V. Parr et al., “Association of Selected Viniviticultural Factors with Sensory and Chemical Characteristics of New Zealand Sauvignon Blanc Wines,” Food Research International 53 (2013): 464—475.

218 trucking the grapes: Dimitra L. Capone and David W. Jeffery, “Effects of Transporting and Processing Sauvignon Blanc Grapes on 3-Mercaptohexan-1-ol Precursor Concentrations,” Journal of Agricultural and Food Chemistry 59 (2011): 4659—4667.

218 unique microbial ecosystem: Nicholas A. Bokulich et al., “Microbial Biogeography of Wine Grapes Is Conditioned by Cultivar, Vintage, and Climate,” Proceedings of the National Academy of Sciences 111 (2014): E139—E148, doi:10.1073/pnas.1317377110.

219 detectably different aroma profile: Sarah Knight et al., “Regional Microbial Signatures Positively Correlate with Differential Wine Phenotypes: Evidence for a Microbial Aspect to Terroir,” Scientific Reports 5 (2015): 14233, doi:10.1038/srep14233.

221 some store better than others: Luke Bell et al., “Use of TD-GC-TOF-MS to Assess Volatile Composition during Post-Harvest Storage in Seven Accessions of Rocket Salad (Eruca sativa),” Food Chemistry 194 (2016): 626—636.

222 vanished after a week: Fernando Vallejo, Francisco Tomás-Barberán, and Cristina García-Viguera, “Health-Promoting Compounds in Broccoli as Influenced by Refrigerated Transport and Retail Sale Period,” Journal of Agricultural and Food Chemistry 51 (2003): 3029—3034.

222 haven’t reached consensus: See, for example, Marcin Baranski et al., “Higher Antioxidant and Lower Cadmium Concentrations and Lower Incidence of Pesticide Residues in Organically Grown Crops: A Systematic Literature Review and Meta-Analyses,” British Journal of Nutrition 112 (2014): 794—811; Diane Bourn and John Prescott, “A Comparison of the Nutritional Value, Sensory Qualities, and Food Safety of Organically and Conventionally Produced Foods,” Critical Reviews in Food Science and Nutrition 42 (2002): 1—34; Alan D. Dangour et al., “Nutritional Quality of Organic Foods: A Systematic Review,” American Journal of Clinical Nutrition 90 (2009): 680—685; Crystal Smith-Spangler et al., “Are Organic Foods Safer or Healthier Than Conventional Alternatives? A Systematic Review,” Annals of Internal Medicine 157 (2012): 348—366.

223 local might not mean fresher: I owe this idea to Alyson Mitchell of the University of California, Davis.

223 didn’t matter one bit: Xin Zhao et al., “Consumer Sensory Analysis of Organically and Conventionally Grown Vegetables,” Journal of Food Science 72 (2007): S87—S91.

224 thought the eco-friendly coffee: Patrik Sörqvist et al., “Who Needs Cream and Sugar When There Is Eco-Labeling? Taste and Willingness to Pay for ’Eco-Friendly’ Coffee,” PLoS One 8 (2013): e80719, doi:10.1371/journal.pone.0080719.

226 says one tomato grower: Quoted in Dan Charles, “How the Taste of Tomatoes Went Bad (and Kept on Going),” NPR All Things Considered, June 28, 2012, accessed March 1, 2016,

227 20 percent less sugar: Ann L. T. Powell et al., “Uniform ripening Encodes a Golden 2-like Transcription Factor Regulating Tomato Fruit Chloroplast Development,” Science 336 (2012): 1711—1715.


233 more of the flavor in the vegetable: Royal Society of Chemistry, “Kitchen Chemistry: The Chemistry of Flavour,”

234 Researchers in England: D. S. Mottram and R. A. Edwards, “The Role of Triglycerides and Phospholipids in the Aroma of Cooked Beef,” Journal of the Science of Food and Agriculture 34 (1983): 517—522.

235 skatole: Peter K. Watkins et al., “Sheepmeat Flavor and the Effect of Different Feeding Systems: A Review,” Journal of Agricultural and Food Chemistry 61 (2013): 3561—3579.

237 621 different Maillard products: Donald S. Mottram and J. Stephen Elmore, “Control of the Maillard Reaction during the Cooking of Food,” in Donald S. Mottram and Andrew J. Taylor, eds., Controlling Maillard Pathways to Generate Flavors (Washington, DC: American Chemical Society Symposium Series 1042, 2010), 143—155.

239 lab version of a cook-off: Chris Kerth, “Determination of Volatile Aroma Compounds in Beef Using Differences in Steak Thickness and Cook Surface Temperature,” Meat Science 117 (2016): 27—35.

240 different sets of microbes: My discussion of cheese styles follows Julie E. Button and Rachel J. Dutton, “Cheese Microbes,” Current Biology 22 (2012): R587—R589.

244 more popular than you’d expect: Michael A. Nestrud, John M. Ennis, and Harry T. Lawless, “A Group-Level Validation of the Supercombinatoriality Property: Finding High-Quality Ingredient Combinations Using Pairwise Information,” Food Quality and Preference 25 (2012): 23—28.

247 experimenting with salty ingredients: Heston Blumenthal, “Weird but Wonderful,” The Guardian, May 4, 2002.

248 Chartier’s book: François Chartier, Taste Buds and Molecules (Hoboken, NJ: Wiley, 2012).

250 studied their molecular overlaps: Yong-Yeol Ahn et al., “Flavor Network and the Principles of Food Pairing,” Scientific Reports 1 (2011): 196, doi:10.1038/srep00196.

251 The Flavor Bible: Karen Page and Andrew Dornenburg, The Flavor Bible: The Essential Guide to Culinary Creativity, Based on the Wisdom of America’s Most Imaginative Chefs (New York: Little, Brown, 2008).

257 a Wassily Kandinsky painting: Charles Michel et al., “A Taste of Kandinsky: Assessing the Influence of the Artistic Visual Presentation of Food on the Dining Experience,” Flavour 3 (2014): 7, doi:10.1186/2044-7248-3-7.

259a quick run through the blender”: Nathan Myhrvold with Chris Young and Maxime Bilet, Modernist Cuisine: The Art and Science of Cooking, Volume 4 (Bellevue, WA: The Cooking Lab, 2011), 343.


261 craft breweries increasing by nearly 20 percent: Brewer’s Association, “Number of Breweries and Brewpubs in U.S.,” accessed May 28, 2016,

261 more than doubling since 2000: Rebecca Smithers, “Good Beer Guide 2015 Shows UK Has Most Breweries per Head of Population,” The Guardian, September 11, 2014,

262 Crisco white sauce: Leslie Brenner, American Appetite: The Coming of Age of a Cuisine (New York: Avon, 1999), 21.

264 The number of farmers’ markets: Anonymous, “Number of US Farmers’ Markets Continues to Rise,” accessed May 28, 2016,

268 found no difference: Wendy V. Parr, David Heatherbell, and K. Geoffrey White, “Demystifying Wine Expertise: Olfactory Threshold, Perceptual Skill and Semantic Memory in Expert and Novice Wine Judges,” Chemical Senses 27 (2002): 747—755.

268 slightly more likely: Gary J. Pickering, Arun K. Jain, and Ram Bezawada, “Super-Tasting Gastronomes? Taste Phenotype Characterization of Foodies and Wine Experts,” Food Quality and Preference 28 (2013): 85—91.