, Volume 223, Issue 3, pp 319–330 | Cite as

Blueberry supplementation induces spatial memory improvements and region-specific regulation of hippocampal BDNF mRNA expression in young rats

  • Catarina Rendeiro
  • David Vauzour
  • Rebecca J. Kean
  • Laurie T. Butler
  • Marcus Rattray
  • Jeremy P. E. Spencer
  • Claire M. WilliamsEmail author
Original Investigation



Flavonoid-rich foods have been shown to be able to reverse age-related cognitive deficits in memory and learning in both animals and humans. However, to date, there have been only a limited number of studies investigating the effects of flavonoid-rich foods on cognition in young/healthy animals.


The aim of this study was to investigate the effects of a blueberry-rich diet in young animals using a spatial working memory paradigm, the delayed non-match task, using an eight-arm radial maze. Furthermore, the mechanisms underlying such behavioural effects were investigated.


We show that a 7-week supplementation with a blueberry diet (2 % w/w) improves the spatial memory performance of young rats (2 months old). Blueberry-fed animals also exhibited a faster rate of learning compared to those on the control diet. These behavioural outputs were accompanied by the activation of extracellular signal-related kinase (ERK1/2), increases in total cAMP-response element-binding protein (CREB) and elevated levels of pro- and mature brain-derived neurotrophic factor (BDNF) in the hippocampus. Changes in hippocampal CREB correlated well with memory performance. Further regional analysis of BDNF gene expression in the hippocampus revealed a specific increase in BDNF mRNA in the dentate gyrus and CA1 areas of hippocampi of blueberry-fed animals.


The present study suggests that consumption of flavonoid-rich blueberries has a positive impact on spatial learning performance in young healthy animals, and these improvements are linked to the activation of ERK–CREB–BDNF pathway in the hippocampus.


Flavonoid Blueberry Learning Memory Brain-derived neurotrophic factor MAP kinase CREB Hippocampus 



This research was supported by the Biotechnology and Biological Sciences Research Council (grant: BB/F008953/1) and the Fundação para a Ciência e a Tecnologia (grant: SFRH/BD/69711/2010) and they were greatly appreciated.


  1. Aberg MA, Aberg ND, Hedbacker H, Oscarsson J, Eriksson PS (2000) Peripheral infusion of IGF-I selectively induces neurogenesis in the adult rat hippocampus. J Neurosci 20:2896–2903PubMedGoogle Scholar
  2. Andres-Lacueva C, Shukitt-Hale B, Galli RL, Jauregui O, Lamuela-Raventos RM, Joseph JA (2005) Anthocyanins in aged blueberry-fed rats are found centrally and may enhance memory. Nutr Neurosci 8:111–120PubMedCrossRefGoogle Scholar
  3. Banerjee S, Smallwood A, Chambers AE, Nicolaides K (2003) Quantitative recovery of immunoreactive proteins from clinical samples following RNA and DNA isolation. Biotechniques 35:450–456PubMedGoogle Scholar
  4. Beking K, Vieira A (2011) Flavonoid intake and disability-adjusted life years due to Alzheimer’s and related dementias: a population-based study involving twenty-three developed countries. Public Health Nutr 13:1403–1409CrossRefGoogle Scholar
  5. Bekinschtein P, Cammarota M, Igaz LM, Bevilaqua LR, Izquierdo I, Medina JH (2007) Persistence of long-term memory storage requires a late protein synthesis- and BDNF-dependent phase in the hippocampus. Neuron 53:261–277PubMedCrossRefGoogle Scholar
  6. Bhutada P, Mundhada Y, Bansod K, Ubgade A, Quazi M, Umathe S, Mundhada D (2010) Reversal by quercetin of corticotrophin releasing factor induced anxiety- and depression-like effect in mice. Prog Neuropsychopharmacol Biol Psychiatry 34:955–960PubMedCrossRefGoogle Scholar
  7. Bito H (1998) A potential mechanism for long-term memory: CREB signaling between the synapse and the nucleus. Seikagaku 70:466–471PubMedGoogle Scholar
  8. Brightwell JJ, Smith CA, Neve RL, Colombo PJ (2007) Long-term memory for place learning is facilitated by expression of cAMP response element-binding protein in the dorsal hippocampus. Learn Mem 14:195–199PubMedCrossRefGoogle Scholar
  9. Casadesus G, Shukitt-Hale B, Stellwagen HM, Zhu X, Lee HG, Smith MA, Joseph JA (2004) Modulation of hippocampal plasticity and cognitive behavior by short-term blueberry supplementation in aged rats. Nutr Neurosci 7:309–316PubMedCrossRefGoogle Scholar
  10. Chan YC, Hosoda K, Tsai CJ, Yamamoto S, Wang MF (2006) Favorable effects of tea on reducing the cognitive deficits and brain morphological changes in senescence-accelerated mice. J Nutr Sci Vitaminol (Tokyo). 52(4):266–73Google Scholar
  11. Choy KH, de Visser Y, Nichols NR, van den Buuse M (2008) Combined neonatal stress and young–adult glucocorticoid stimulation in rats reduce BDNF expression in hippocampus: effects on learning and memory. Hippocampus 18:655–667PubMedCrossRefGoogle Scholar
  12. Cohen-Cory S, Fraser SE (1995) Effects of brain-derived neurotrophic factor on optic axon branching and remodelling in vivo. Nature 378:192–196PubMedCrossRefGoogle Scholar
  13. Conkright MD, Guzman E, Flechner L, Su AI, Hogenesch JB, Montminy M (2003) Genome-wide analysis of CREB target genes reveals a core promoter requirement for cAMP responsiveness. Mol Cell 11:1101–1108PubMedCrossRefGoogle Scholar
  14. Conner JM, Lauterborn JC, Yan Q, Gall CM, Varon S (1997) Distribution of brain-derived neurotrophic factor (BDNF) protein and mRNA in the normal adult rat CNS: evidence for anterograde axonal transport. J Neurosci 17:2295–2313PubMedGoogle Scholar
  15. Countryman RA, Gold PE (2007) Rapid forgetting of social transmission of food preferences in aged rats: relationship to hippocampal CREB activation. Learn Mem 14:350–358PubMedCrossRefGoogle Scholar
  16. Cunha C, Brambilla R, Thomas KL (2010) A simple role for BDNF in learning and memory? Front Mol Neurosci 3:1PubMedGoogle Scholar
  17. Dinges DF (2006) Cocoa flavanols, cerebral blood flow, cognition, and health: going forward. J Cardiovasc Pharmacol 47(Suppl 2):S221–S223PubMedGoogle Scholar
  18. Duffy KB, Spangler EL, Devan BD, Guo Z, Bowker JL, Janas AM, Hagepanos A, Minor RK, Decabo R, Mouton PR, Shukitt-Hale B, Joseph JA, Ingram DK (2007) A blueberry-enriched diet provides cellular protection against oxidative stress and reduces a kainate-induced learning impairment in rats. Neurobiol AgingGoogle Scholar
  19. Falkenberg T, Mohammed AK, Henriksson B, Persson H, Winblad B, Lindefors N (1992) Increased expression of brain-derived neurotrophic factor mRNA in rat hippocampus is associated with improved spatial memory and enriched environment. Neurosci Lett 138:153–156PubMedCrossRefGoogle Scholar
  20. Fisher ND, Sorond FA, Hollenberg NK (2006) Cocoa flavanols and brain perfusion. J Cardiovasc Pharmacol 47(Suppl 2):S210–S214PubMedCrossRefGoogle Scholar
  21. Francis ST, Head K, Morris PG, Macdonald IA (2006) The effect of flavanol-rich cocoa on the fMRI response to a cognitive task in healthy young people. J Cardiovasc Pharmacol 47(Suppl 2):S215–S220PubMedCrossRefGoogle Scholar
  22. Gomez-Pinilla F (2008) Brain foods: the effects of nutrients on brain function. Nat Rev Neurosci 9:568–578PubMedCrossRefGoogle Scholar
  23. Goyarzu P, Malin DH, Lau FC, Taglialatela G, Moon WD, Jennings R, Moy E, Moy D, Lippold S, Shukitt-Hale B, Joseph JA (2004) Blueberry supplemented diet: effects on object recognition memory and nuclear factor-kappa B levels in aged rats. Nutr Neurosci 7:75–83PubMedCrossRefGoogle Scholar
  24. Haque AM, Hashimoto M, Katakura M, Tanabe Y, Hara Y, Shido O (2006) Long-term administration of green tea catechins improves spatial cognition learning ability in rats. J Nutr 136:1043–1047PubMedGoogle Scholar
  25. Hartman RE, Shah A, Fagan AM, Schwetye KE, Parsadanian M, Schulman RN, Finn MB, Holtzman DM (2006) Pomegranate juice decreases amyloid load and improves behavior in a mouse model of Alzheimer’s disease. Neurobiol Dis 24(3):506–15Google Scholar
  26. Hoffman JR, Donato A, Robbins SJ (2004) Ginkgo biloba promotes short-term retention of spatial memory in rats. Pharmacol Biochem Behav 77:533–539PubMedCrossRefGoogle Scholar
  27. Hou Y, Aboukhatwa MA, Lei DL, Manaye K, Khan I, Luo Y (2010) Anti-depressant natural flavonols modulate BDNF and beta amyloid in neurons and hippocampus of double TgAD mice. Neuropharmacology 58:911–920PubMedCrossRefGoogle Scholar
  28. Jeon SJ, Rhee SY, Seo JE, Bak HR, Lee SH, Ryu JH, Cheong JH, Shin CY, Kim GH, Lee YS, Ko KH (2011) Oroxylin A increases BDNF production by activation of MAPK-CREB pathway in rat primary cortical neuronal culture. Neurosci Res 69:214–222PubMedCrossRefGoogle Scholar
  29. Jerman T, Kesner RP (2006) Disconnection analysis of CA3 and DG in mediating encoding but not retrieval in a spatial maze learning task. Learn Mem 13:458–464Google Scholar
  30. Joseph JA, Shukitt-Hale B, Denisova NA, Prior RL, Cao G, Martin A, Taglialatela G, Bickford PC (1998) Long-term dietary strawberry, spinach, or vitamin E supplementation retards the onset of age-related neuronal signal-transduction and cognitive behavioral deficits. J Neurosci 18:8047–8055PubMedGoogle Scholar
  31. Joseph JA, Shukitt-Hale B, Denisova NA, Bielinski D, Martin A, McEwen JJ, Bickford PC (1999) Reversals of age-related declines in neuronal signal transduction, cognitive, and motor behavioral deficits with blueberry, spinach, or strawberry dietary supplementation. J Neurosci 19:8114–8121PubMedGoogle Scholar
  32. Joseph JA, Denisova NA, Arendash G, Gordon M, Diamond D, Shukitt-Hale B, Morgan D (2003) Blueberry supplementation enhances signaling and prevents behavioral deficits in an Alzheimer disease model. Nutr Neurosci 6:153–162PubMedCrossRefGoogle Scholar
  33. Kaur T, Pathak CM, Pandhi P, Khanduja KL (2008) Effects of green tea extract on learning, memory, behavior and acetylcholinesterase activity in young and old male rats. Brain Cogn 67(1):25–30Google Scholar
  34. Kelleher RJ 3rd, Govindarajan A, Tonegawa S (2004) Translational regulatory mechanisms in persistent forms of synaptic plasticity. Neuron 44:59–73PubMedCrossRefGoogle Scholar
  35. Krikorian R, Shidler MD, Nash TA, Kalt W, Vinqvist-Tymchuk MR, Shukitt-Hale B, Joseph JA (2010) Blueberry supplementation improves memory in older adults. J Agric Food Chem 58:3996–4000PubMedCrossRefGoogle Scholar
  36. Kromhout D, Menotti A, Kesteloot H, Sans S (2002) Prevention of coronary heart disease by diet and lifestyle: evidence from prospective cross-cultural, cohort, and intervention studies. Circulation 105:893–898PubMedCrossRefGoogle Scholar
  37. Kuriyama S, Hozawa A, Ohmori K, Shimazu T, Matsui T, Ebihara S, Awata S, Nagatomi R, Arai H, Tsuji I (2006) Green tea consumption and cognitive function: a cross-sectional study from the Tsurugaya Project 1. Am J Clin Nutr 83(2):355–361Google Scholar
  38. Lai HC, Chao WT, Chen YT, Yang VC (2004) Effect of EGCG, a major component of green tea, on the expression of Ets-1, c-Fos, and c-Jun during angiogenesis in vitro. Cancer Lett 213(2):181–188Google Scholar
  39. Lassalle JM, Bataille T, Halley H (2000) Reversible inactivation of the hippocampal mossy fiber synapses in mice impairs spatial learning, but neither consolidation nor memory retrieval, in the Morris navigation task. Neurobiology of learning and memory 73(3):243–257Google Scholar
  40. Lee I, Kesner RP (2003) Differential roles of dorsal hippocampal subregions in spatial working memory with short versus intermediate delay. Behav Neurosci 117:1044–1053PubMedCrossRefGoogle Scholar
  41. Lee I, Kesner RP (2004) Encoding versus retrieval of spatial memory: double dissociation between the dentate gyrus and the perforant path inputs into CA3 in the dorsal hippocampus. Hippocampus 14(1):66–76Google Scholar
  42. Letenneur L, Proust-Lima C, Le Gouge A, Dartigues JF, Barberger-Gateau P (2007) Flavonoid intake and cognitive decline over a 10-year period. Am J Epidemiol 165:1364–1371PubMedCrossRefGoogle Scholar
  43. Leutgeb S, Leutgeb JK, Treves A, Moser MB, Moser EI (2004) Distinct ensemble codes in hippocampal areas CA3 and CA1. Science 305:1295–1298PubMedCrossRefGoogle Scholar
  44. Li Q, Zhao HF, Zhang ZF, Liu ZG, Pei XR, Wang JB, Cai MY, Li Y (2009a) Long-term administration of green tea catechins prevents age-related spatial learning and memory decline in C57BL/6 J mice by regulating hippocampal cyclic amp-response element binding protein signaling cascade. Neuroscience 159:1208–1215PubMedCrossRefGoogle Scholar
  45. Li Q, Zhao HF, Zhang ZF, Liu ZG, Pei XR, Wang JB, Li Y (2009b) Long-term green tea catechin administration prevents spatial learning and memory impairment in senescence-accelerated mouse prone-8 mice by decreasing Abeta1-42 oligomers and upregulating synaptic plasticity-related proteins in the hippocampus. Neuroscience 163:741–749PubMedCrossRefGoogle Scholar
  46. Liu RY, Fioravante D, Shah S, Byrne JH (2008) cAMP response element-binding protein 1 feedback loop is necessary for consolidation of long-term synaptic facilitation in Aplysia. J Neurosci 28:1970–1976PubMedCrossRefGoogle Scholar
  47. Lowry OH, Rosebrough NJ, Farr AL, Randall RJ (1951) Protein measurement with the Folin phenol reagent. J Biol Chem 193:265–275PubMedGoogle Scholar
  48. Macready AL, Kennedy OB, Ellis JA, Williams CM, Spencer JP, Butler LT (2009) Flavonoids and cognitive function: a review of human randomized controlled trial studies and recommendations for future studies. Genes Nutr 4:227–242PubMedCrossRefGoogle Scholar
  49. McAllister AK, Katz LC, Lo DC (1999) Neurotrophins and synaptic plasticity. Annu Rev Neurosci 22:295–318PubMedCrossRefGoogle Scholar
  50. Mizuno M, Yamada K, Olariu A, Nawa H, Nabeshima T (2000) Involvement of brain-derived neurotrophic factor in spatial memory formation and maintenance in a radial arm maze test in rats. J Neurosci 20:7116–7121PubMedGoogle Scholar
  51. Mohamed HA, Yao W, Fioravante D, Smolen PD, Byrne JH (2005) cAMP-response elements in Aplysia creb1, creb2, and Ap-uch promoters: implications for feedback loops modulating long-term memory. J Biol Chem 280:27035–27043PubMedCrossRefGoogle Scholar
  52. Nakamura EM, da Silva EA, Concilio GV, Wilkinson DA, Masur J (1991) Reversible effects of acute and long-term administration of delta-9-tetrahydrocannabinol (THC) on memory in the rat. Drug Alcohol Depend 28:167–175PubMedCrossRefGoogle Scholar
  53. Oliveira DR, Sanada PF, Saragossa Filho AC, Innocenti LR, Oler G, Cerutti JM, Cerutti SM (2009) Neuromodulatory property of standardized extract Ginkgo biloba L. (EGb 761) on memory: behavioral and molecular evidence. Brain Res 1269:68–89PubMedCrossRefGoogle Scholar
  54. Olton DS, Papas BC (1979) Spatial memory and hippocampal function. Neuropsychologia 17:669–682PubMedCrossRefGoogle Scholar
  55. Olton DS, Samuelson RJ (1976) Remembrance of places passed—spatial memory in rats. J Exp Psychol Anim Behav Process 2:97–116CrossRefGoogle Scholar
  56. Parrott MD, Greenwood CE (2007) Dietary influences on cognitive function with aging: from high-fat diets to healthful eating. Ann N Y Acad Sci 1114:389–397PubMedCrossRefGoogle Scholar
  57. Phillips HS, Hains JM, Laramee GR, Rosenthal A, Winslow JW (1990) Widespread expression of BDNF but not NT3 by target areas of basal forebrain cholinergic neurons. Science 250:290–294PubMedCrossRefGoogle Scholar
  58. Ramirez MR, Izquierdo I, do Carmo Bassols Raseira M, Zuanazzi JA, Barros D, Henriques AT (2005) Effect of lyophilised Vaccinium berries on memory, anxiety and locomotion in adult rats. Pharmacol Res 52:457–462PubMedCrossRefGoogle Scholar
  59. Rattray M, Michael GJ, Lee J, Wotherspoon G, Bendotti C, Priestley JV (1999) Intraregional variation in expression of serotonin transporter messenger RNA by 5-hydroxytryptamine neurons. Neuroscience 88:169–183PubMedCrossRefGoogle Scholar
  60. Rendeiro C, Spencer JP, Vauzour D, Butler LT, Ellis JA, Williams CM (2009) The impact of flavonoids on spatial memory in rodents: from behaviour to underlying hippocampal mechanisms. Genes Nutr 4(4):251–270PubMedCrossRefGoogle Scholar
  61. Ried K, Sullivan T, Fakler P, Frank OR, Stocks NP (2010) Does chocolate reduce blood pressure? A meta-analysis. BMC Med 8:39PubMedCrossRefGoogle Scholar
  62. Robbins RJ, Leonczak J, Johnson JC, Li J, Kwik-Uribe C, Prior RL, Gu L (2009) Method performance and multi-laboratory assessment of a normal phase high pressure liquid chromatography-fluorescence detection method for the quantitation of flavanols and procyanidins in cocoa and chocolate containing samples. J Chromatogr A 1216:4831–4840PubMedCrossRefGoogle Scholar
  63. Rolls ET, Kesner RP (2006) A computational theory of hippocampal function, and empirical tests of the theory. Prog Neurobiol 79:1–48PubMedCrossRefGoogle Scholar
  64. Schaaf MJ, Workel JO, Lesscher HM, Vreugdenhil E, Oitzl MS, de Kloet ER (2001) Correlation between hippocampal BDNF mRNA expression and memory performance in senescent rats. Brain Res 915:227–233PubMedCrossRefGoogle Scholar
  65. Schroeter H, Bahia P, Spencer JP, Sheppard O, Rattray M, Cadenas E, Rice-Evans C, Williams RJ (2007) (−)Epicatechin stimulates ERK-dependent cyclic AMP response element activity and up-regulates GluR2 in cortical neurons. J Neurochem 101:1596–1606PubMedCrossRefGoogle Scholar
  66. Sekeres MJ, Neve RL, Frankland PW, Josselyn SA (2010) Dorsal hippocampal CREB is both necessary and sufficient for spatial memory. Learn Mem 17:280–283PubMedCrossRefGoogle Scholar
  67. Shif O, Gillette K, Damkaoutis CM, Carrano C, Robbins SJ, Hoffman JR (2006) Effects of Ginkgo biloba administered after spatial learning on water maze and radial arm maze performance in young adult rats. Pharmacol Biochem Behav 84:17–25PubMedCrossRefGoogle Scholar
  68. Shukitt-Hale B, Lau FC, Carey AN, Galli RL, Spangler EL, Ingram DK, Joseph JA (2008) Blueberry polyphenols attenuate kainic acid-induced decrements in cognition and alter inflammatory gene expression in rat hippocampus. Nutr Neurosci 11:172–182PubMedCrossRefGoogle Scholar
  69. Silhol M, Arancibia S, Maurice T, Tapia-Arancibia L (2007) Spatial memory training modifies the expression of brain-derived neurotrophic factor tyrosine kinase receptors in young and aged rats. Neuroscience 146:962–973PubMedCrossRefGoogle Scholar
  70. Spencer JP (2008) Flavonoids: modulators of brain function? Br J Nutr 99 E Suppl 1:ES60–77Google Scholar
  71. Sweatt JD (2001) The neuronal MAP kinase cascade: a biochemical signal integration system subserving synaptic plasticity and memory. J Neurochem 76:1–10PubMedCrossRefGoogle Scholar
  72. Sweatt JD (2004) Mitogen-activated protein kinases in synaptic plasticity and memory. Curr Opin Neurobiol 14:311–317PubMedCrossRefGoogle Scholar
  73. Tao X, Finkbeiner S, Arnold DB, Shaywitz AJ, Greenberg ME (1998) Ca2+ influx regulates BDNF transcription by a CREB family transcription factor-dependent mechanism. Neuron 20:709–726PubMedCrossRefGoogle Scholar
  74. Unno K, Takabayashi F, Yoshida H, Choba D, Fukutomi R, Kikunaga N, Kishido T, Oku N, Hoshino M (2007) Daily consumption of green tea catechin delays memory regression in aged mice. Biogerontology 8(2):89–95Google Scholar
  75. Vago DR, Kesner RP (2005) An electrophysiological and behavioural characterization of the temporoammonic pathway: disruption produces deficits in retrieval and spatial mismatch. Soc Neurosci Abstr 647:5Google Scholar
  76. van Praag H (2008) Neurogenesis and exercise: past and future directions. Neuromolecular Med 10:128–140PubMedCrossRefGoogle Scholar
  77. van Praag H (2009) Exercise and the brain: something to chew on. Trends Neurosci 32:283–290PubMedCrossRefGoogle Scholar
  78. van Praag H, Lucero MJ, Yeo GW, Stecker K, Heivand N, Zhao C, Yip E, Afanador M, Schroeter H, Hammerstone J, Gage FH (2007) Plant-derived flavanol (−)epicatechin enhances angiogenesis and retention of spatial memory in mice. J Neurosci 27:5869–5878PubMedCrossRefGoogle Scholar
  79. Vauzour D, Vafeiadou K, Rice-Evans C, Williams RJ, Spencer JP (2007) Activation of pro-survival Akt and ERK1/2 signalling pathways underlie the anti-apoptotic effects of flavanones in cortical neurons. J Neurochem 103(4):1355–1367Google Scholar
  80. Vignes M, Maurice T, Lante F, Nedjar M, Thethi K, Guiramand J, Recasens M (2006) Anxiolytic properties of green tea polyphenol (−)-epigallocatechin gallate (EGCG). Brain Res 1110:102–115PubMedCrossRefGoogle Scholar
  81. Wang Y, Wang L, Wu J, Cai J (2006) The in vivo synaptic plasticity mechanism of EGb 761-induced enhancement of spatial learning and memory in aged rats. Br J Pharmacol 148:147–153PubMedCrossRefGoogle Scholar
  82. Ward CP, Redd K, Williams BM, Caler JR, Luo Y, McCoy JG (2002) Ginkgo biloba extract: cognitive enhancer or antistress buffer. Pharmacol Biochem Behav 72:913–922PubMedCrossRefGoogle Scholar
  83. Williams CM, El Mohsen MA, Vauzour D, Rendeiro C, Butler LT, Ellis JA, Whiteman M, Spencer JP (2008) Blueberry-induced changes in spatial working memory correlate with changes in hippocampal CREB phosphorylation and brain-derived neurotrophic factor (BDNF) levels. Free Radic Biol Med 45:295–305PubMedCrossRefGoogle Scholar
  84. Williams RJ, Spencer JP, Rice-Evans C (2004) Flavonoids: antioxidants or signalling molecules? Free Radic Biol Med 36(7):838–49Google Scholar
  85. Zhao C, Deng W, Gage FH (2008) Mechanisms and functional implications of adult neurogenesis. Cell 132:645–660PubMedCrossRefGoogle Scholar

Copyright information

© Springer-Verlag 2012

Authors and Affiliations

  • Catarina Rendeiro
    • 1
    • 3
  • David Vauzour
    • 2
  • Rebecca J. Kean
    • 3
  • Laurie T. Butler
    • 3
  • Marcus Rattray
    • 4
  • Jeremy P. E. Spencer
    • 1
  • Claire M. Williams
    • 3
    Email author
  1. 1.Molecular Nutrition Group, School of Chemistry, Food and PharmacyUniversity of ReadingReadingUK
  2. 2.Department of Nutrition, Norwich Medical School, Faculty of Medicine and Health SciencesUniversity of East AngliaNorwichUK
  3. 3.School of Psychology and Clinical Language SciencesUniversity of ReadingReadingUK
  4. 4.Reading School of PharmacyUniversity of ReadingReadingUK

Personalised recommendations