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Preview REVIEW Effect of sensory perception of foods on appetite and food intake

InternationalJournalofObesity(2003)27,1152–1166 &2003NaturePublishingGroup Allrightsreserved 0307-0565/03 $25.00 www.nature.com/ijo REVIEW Effect of sensory perception of foods on appetite and food intake: a review of studies on humans LB S(cid:1)rensen1*, P M(cid:1)ller2, A Flint1, M Martens2 and A Raben1 1Department of Human Nutrition, Centre for Advanced Food Studies, The Royal Veterinary and Agricultural University, Frederiksberg C, Denmark; and 2Department of Dairy and Food Science, The Royal Veterinary and Agricultural University, Frederiksberg C, Denmark Objective:Howmuchdothesensorypropertiesoffoodinfluencethewaypeopleselecttheirfoodandhowmuchtheyeat?The objectiveofthispaperistoreviewresultsfromstudiesinvestigatingthelinkbetweenthesensoryperceptionoffoodandhuman appetite regulation. Content of the review: The influence of palatability on appetite and food intake in humans has been investigated in several studies. All reviewed studies have shown increased intake as palatability increased, whereas assessments of the effect of palatabilityusingmeasuresofsubjectiveappetitesensationshaveshowndivergingresults,forexample,subjectseitherfeelmore hungryandlessfullafterapalatablemealcomparedtoalesspalatablemeal,ortheyfeeltheopposite,orthereisnodifference. Whether palatability hasaneffecton appetite intheperiod following consumption of atest meal isunclear. Severalstudieshaveinvestigatedwhichsensorypropertiesoffoodareinvolvedinsensory-specificsatiety.Taste,smell,texture andappearance-specificsatietieshavebeenidentified,whereasstudiesontheroleofmacronutrientsandtheenergycontentof thefoodinsensory-specificsatietyhavegivenequivocalresults.Differentstudieshaveshownthatmacronutrientsandenergy content play a role in sensory-specific satiety or that macronutrients and energy content are not a factor in sensory-specific satiety. Sensory-specific satiety may have an important influence on the amount of food eaten. Studies have shown that increasingthefoodvarietycanincreasefoodandenergyintakeandintheshorttomediumtermalterenergybalance.Further knowledgeabouttheimportanceofflavourinappetiteregulationisneeded,forexample,whichflavourcombinationsimprove satietymost,thepossibleconnectionbetweenflavourintensityandsatiety,theeffectofpersistenceofchemestheticsensation onpalatabilityandsatiety,andtowhatextentgeneticvariationintastesensitivityandperceptioninfluencesdietaryhabitsand weightcontrol. International Journal ofObesity (2003) 27, 1152–1166. doi:10.1038/sj.ijo.0802391 Keywords: appetite;hunger; food intake; palatability; sensory-specific satiety; intensity Introduction impression people get from the sensory properties of foods How much do the sensory properties of food influence playsaveryimportantroleinthewaytheyselecttheirfood eating behaviour? Traditionally, sensory science and nutri- andhowmuchtheyeat.Seeninthelightofthecontinuing tion,includingtheuncoveringofthemechanismsofhuman increaseintheprevalenceofoverweightandobesityinlarge appetite regulation, have been two separate fields of parts of the world,1 it isimportant to identify all aspects of research. However, within the last decades, there has been humanappetiteregulationtounderstandthistrend,andto increasing interaction between the two fields, which has preventitfromprogressing.Thus,theobjectiveofthispaper resultedinawiderangeofstudiesthathavecoveredaspects is to review results from studies investigating the link frombothfields.Resultsfromthesestudiessuggestthatthe between the sensory perception of food and human eating behaviour. The review begins with a presentation of how appetiteandpalatabilityaremeasured. *Correspondence:LBSørensen,DepartmentofHumanNutrition,Centre for Advanced Food Studies, The Royal Veterinary and Agricultural University,Rolighedsvej30,DK-1958FrederiksbergC,Denmark. E-mail:[email protected] Methods ThestudywassupportedbyOrklaFoodsAS,theDanishDairyResearch Intheselectionofstudies,wehaveconcentratedonhuman Foundation(DanishDairyBoard)andtheDanishGovernment. studies with participation of healthy subjects with a body Received1July2002;revised11April2003; accepted7May2003 massindexof18.5–25.0kg/m2.Normal-weightsubjectswere Sensoryscienceandappetite LBSørensenetal 1153 chosen because it is important to identify how appetite is manipulations.Thisismadepossiblebyservinganadlibitum regulatedinnormal-weightpeopleinordertoprovideabasis meal and asking the subjects to eat until they are full and for understanding whether there is a difference in normal- then recording the amount of food eaten in grams or weight and overweight people’s eating behaviour. The units.11,12 Stubbs et al13 have reviewed the use of VAS as a subjects were adults (18–50y) and the women were pre- tool for measuring subjective appetite sensations and they menopausal. Only studies conducted in Western countries concluded that VAS show some ability to predict aspects of published in 1980 and onwards were included. Selected feedingbehaviourandactasausefuladjuncttomeasuresof studies had a link between the sensory perception of food food, energy and nutrient intake; VAS show sensitivity to and appetite sensations and/or energy intake. Therefore, experimentalmanipulations andshowgoodreproducibility studies dealing only with macronutrients and appetite provided that they are used in within-subject designs. Flint sensations and/or energy intake were not included. In the etal14havestudiedthereproducibility,powerandvalidityof search for relevant studies, we used the databases Medline VASandtheyfoundthatusingapaireddesign,whenlooking andFoodScienceandTechnologyAbstractsand,inaddition, for an effect size of 10%, 18 subjects are needed to achieve referencelistswerescannedthoroughly. sufficientstudypowerandtherebyminimisingtheriskofa type IIerror (‘false-negative’ result). Iftheeffect parameters ofinterestarelimitedtomeanpostprandialappetiteratings, Measuringappetiteinhumansubjects eightsubjectsareenough. To prevent confusion about the terminology used when When investigating human appetite regulation using describing aspects of appetite regulation, the following will female subjects, it is very important that the women are in be applied in the review: ‘appetite’ is often referred to as a the same phase in their menstrual cycle on all test days sensationrelatedtomaintenanceofeating,oftenadesirefor becauseoftheuncertaintyofhowthedifferentphasesofthe somethingspecific,2butinthisreviewitisusedasageneral menstrual cycle influence appetite regulation. Dye and term of overall sensations related to food intake. The Blundell15 have reviewed 30 studies from the period 1981 definitionof‘satiety’canbeseparatedintotwofunctionally to 1995 all investigating the connection between the different terms: ‘satiation’ or ‘within-meal’ satiety, which menstrual cycle and energy intake. With the reservation referstotheprocessthatleadstotheterminationofameal, fordifferentstudydesignskeptinmind,thegeneralpicture and‘satiety’or‘between-mealsatiety’,whichisconsideredas shows that women have asignificantly higher spontaneous the state of inhibition of eating, and therefore related to energy intake in the premenstrual phase (luteal phase) the time interval until the next eating episode.3–5 ‘Hunger’ comparedtothepostmenstrualphase(follicularphase). canbedescribedasanagging,irritatingfeelingthatsignifies food deprivation to a degree that the next eating episode should take place.2,3,6 The term ‘fullness’ is defined as a Measuringpalatabilityinhumansubjects sensation of the degree of stomach filling, and ‘prospective There have been several suggestions to define palatability, foodconsumption’asanindicatorofthesupposedamount but in his review on palatability and appetite Yeomans16 offorthcomingfoodintake. suggests that palatability should be defined as the hedonic Within appetite research, different scales have been used evaluation of oro-sensory food cues under standardised for the assessment of subjective appetite sensations. Fixed- conditions. The sensory properties of food determining point scales (FPS) and visual analogue scales (VAS) are the palatability are taste, smell, texture, temperature, visual most commonly used rating methods. FPS usually uses appearance, sound and trigeminal input (irritative sensa- numerical scales, which vary greatly in complexity and tions).17 structure.Thus,theintervalsorpointsmayhavenumerical Palatability has been measured using either VAS or FPS indications or different wording.7 Most often used in where subjects are asked questions like ‘how pleasant have appetiteresearchareVAS,intheformofhorizontallinesof you found the food?’, ‘how satisfying have you found the varyinglength,withwordsanchoredateachend,describing food?’, ‘how palatable is this food?’, ‘how attractive do you theextremesofaunipolarquestion(forexampleforhunger: findthefood?’or‘howmuchdidyoulikethefood?’.7,8,18,19 ‘Ihaveneverbeenmorehungry’/’Iamnothungryatall’).8–11 Palatability is classified according to how pleasant or well Subjects are asked to make a vertical mark across the line likedthefoodis,themorepleasantorliked,thehigherthe corresponding to their feeling at the present time. Quanti- palatability. Many investigators have used the terms palat- fication of the measurement is made by measuring the abilityandpleasantnesssynonymously,butthereisadanger distance from the left end of the line to the mark. The in this because of different interpretations of the term subjects are asked about their appetite for a meal or their palatability.11 feelings of hunger, desire to eat, satiety, fullness and pro- The palatability of food is often assessed by taste tests spective food consumption, with text expressing the most pleasantness ratings, where a small sample of food is tasted positiveandmostnegativeratinganchoredateachend. and swallowed,20–23 or by using taste-and-spit pleasantness Besidesusingappetiteratingstoassessappetitesensations, tests,wherethefoodsamplesaretastedandthenspitout.24 food intake has been used to measure the effect of dietary Investigationsofwhetherthesetestscanpredictconsumption InternationalJournalofObesity Sensoryscienceandappetite LBSørensenetal 1154 have shown that taste test pleasantness ratings are poor there were no differences in intake in the semi-liquid and predictors of the actual amount a person will consume in sandwich meals (see Table 1). In this study, appetite was comparisontootherpeople,buttheycanbereliableguides measuredusingFPS,andtheresultsshowednodifferencein toconsumptiondifferenceswithinaperson,25andthattaste- effectofpalatabilityineitherofthemealtypes.Inastudy,by and-spit pleasantness ratings cannot be used to predict Hill et al,8 where the effect of palatability on appetite was consumption by individual subjects, but perhaps can be measuredsolelyusingVAS,adifferenceinratingsofdesireto usedtopredicttheaverageconsumptionofaproduct,26and eat was observed as soon as the subjects saw the food, the theresultsshouldbetakencautiouslywhentryingtopredict difference mostly being due to an increase in ratings in ingestion.27 response to the highly preferred meal. The difference persisted throughout the meal declining only after the mealhadended8(seeTable1).Inaddition,theinvestigators Studies investigating the link between the sensory foundthat2hafterthetestmeal,ratingsofdesiretoeatand properties of food and appetite hunger were significantly higher after consumption of the Influenceofpalatabilityonappetiteandfoodintake preferredfood.8 Theusualapproachinstudyingtheinfluenceofpalatability Sweetness is a majorcontributor to palatability, although on appetite has been to offer subjects test foods with thereareindividualdifferencesinoptimallevelofsweetness. differentlevelsofpalatabilityandcomparetheeffectofthe In two studies, the palatability of the test food was testfoodsonhunger,satietyand/orsubsequentfoodintake. manipulatedbyusingyoghurtwithdifferentconcentrations One way is to make use of subjects’ food preferences by of the intense sweetener, aspartame, and sucrose, respec- making a subjective index of the palatability of test foods tively, hereby making use of the subjects’ preferences for obtained from each subject using VAS and then comparing sweettaste.31,32Inbothstudies,yoghurtintakewasgreaterat this index with the amount eaten by the subjects. Another the most preferred concentration than at all the other way to make use of the subjects’ preferences for foods is by concentrations, and in the study using aspartame, sponta- comparing the effect of a preferred test meal and a neous 24-h food intake was significantly greater following nonpreferred test meal on appetite, the test meals being consumption of the preferred yoghurt, however, only in equicaloricalthoughnotidenticalinmacronutrientcontent. men.31 Studiesusingtheseapproacheshaveshownincreasedintake Speculations as to whether factors other than palatability aspalatabilityincreased12,28–30(seeTable1).Guy-Grandetal7 could explain differences in appetite can be avoided by compared the effect of two levels of palatability in three manipulating the palatability of the test foods by factors different types of test meals (conventional, semiliquid with little effect on the nutritional quality. To manipulate and sandwich), and they observed that high palatability flavourintestmealsaddingmustard,spicessuchasoregano increased intake only in the conventional meal, whereas orcumin,citricacidorsalt,orvaryingthequalityofcheese Table1 Effectofpalatabilityusingpreferredfoods Study Subjects Testmeal Hungerratings Foodintake BellisleandLeMagnen28,29 3males 5differentflavouredsandwiches NR Subjectivepalatabilityscores (1980,1981) 7females plusamixedmealinwhichall correlatedsignificantlywith Withinsubjects 5flavourswereoffered. numberofunitseatenin5 Mealswereequalinenergy subjectsoutof10 Bellisleetal12(1984) 5males Leastormostacceptablesandwich NR Mostacceptablesandwich 5females type(units)identifiedforeachsubject typeincreasedintake Withinsubjects Hilletal8(1984) 12females Ahighlypreferredandaless Increasedafterthesightof NR Withinsubjects preferredmealmadeforeach preferredfood.Increases individual.Mealswere 2haftereatingthepreferredmeal equicaloric Spiegeletal30(1989) 9females High-andlow-palatablefillingin NR Highpalatabilityincreased Withinsubjects spiralsofbread intake Guy-Grandetal7(1994) 7femalesa Conventional,semiliquidand Nodifferences Conventional:highpalatability Withinsubjects sandwichmealsat2levelsof increasedintake.Semiliquid: palatability.Mealswere nodifference.Sandwich: equalinenergy nodifference NR:notreported. aThewomenweretestedbetweenthe10thandthe25thdayoftheirmenstrualcycle. InternationalJournalofObesity Sensoryscienceandappetite LBSørensenetal 1155 Table2 Theeffectofpalatability Study Subjects Preload Testmeal Hungerratings Foodintake Bobroffand 8males None Banana-coladafrozen Morehungryandless Highpalatabilityincreased Kissileff25(1986) 8females yoghurt:plain,6gcumin/ satiatedaftercuminmeals intake Withinsubjects 1500gdrinkor4.5g cumin/1700gdrink Bellisleetal35 36malesand None Spinachmousseorbeef NR Highpalatabilityincreased (1989) females andvegetablejelly:0% intake Betweensubjects monosodiumglutamate (MSG),0.6%MSG,1.2% MSG Rogersand Inallstudies: Inallstudies: Study1:notestmeal Study1:nodifferences Study1:NR Blundell36(1990) 13males Nopreloadorbeef Study2:lunch,30min Study2:NR Study2:noeffect 23females consomme´with0%MSG, delay Study3:nodifferences Study3:noeffect 4subjects 10%MSG(study2only), Study3:lunch,2mindelay afterpreload.No excluded 20%MSG differencesimmediately Withinsubjects Fixedintake aftertestmeal.Duringnext 3hmorerapidrecoveryin hungeranddesiretoeat after20%MSGpreload Warwicketal10 7males Tasty-CHO, Standardlunch,5hdelay Lesshungryandmorefull Noeffect (1993) 3femalesa bland-CHO,tasty-fat, aftertastybreakfast Withinsubjects bland-fat.Fixedintake Yeomans18(1996) 27males None Pastaintomatosauce:no Nodifferenceexceptfor Highpalatabilityincreased 27females oregano:bland;0.27% fullnessratingsreflecting intake Betweensubjects oregano:palatable;0.54% theamounteaten oregano:strong Yeomansetal33 16males None Pastaintomatosauce: Hungerwaslowerinthe Highpalatabilityincreased (1997) Withinsubjects bland,nooregano; continuous/bland.Fullness intake palatable,0.27%oregano. wasunaffectedbytypeof 2eatingstyles: foodeaten continuous/interrupted DeGraafetal19 9males Tomatosoup;nocitricacid: Avarietyoffoodproducts, Lowerhungerratingsafter Highpalatabilityincreased (1999) 29females palatable,0.75gcitric 15or90mindelay thehigherpalatableadlib intakeofadlibpreload Withinsubjects acid/100g:med. preloads. Noeffectontestmeal palatable;1.5gcitricacid/ Withfixedpreloadintake intakeineitherthe15or 100g:unpalatable therewasnoeffectof 90mincondition Adliborfixedintake palatabilityeitherinthe 15or90mincondition Yeomansand 50males None Pastaincheesesauce:bland Nodifferences Highpalatabilityincreased Symes11(1999) Withinsubjects withmildtastingcheese, intake palatablewithamature cheeseversionand mustard Zandstraetal34 11males None Sandwicheswithspread Fortheleastpleasant Palatabilityeffectonthe (2000) 25females madeofbread:0%NaCl: bread,desiretoeatand firstday.Nodifferenceon Withinsubjects leastpleasant;4%NaCl: fullnessincreasedover thefifthday med.pleasant,2%NaCl: days,nodifferencefor mostpleasant.For3 med.andmostpleasant weeksF5dayspersalt bread level NR:notreported. aThewomenweretestedduringthefollicularstageoftheirmenstrualcycle. have all been used.10,11,18,19,25,33,34 In addition, monoso- but one,35 of the studies also investigated the effect of dium glutamate has been used to enhance palatability.35,36 palatabilityonappetiteusingVASandawiderangeofresults All, but two,10,36 of the above-mentioned studies show emerged when reviewing the studies: in Bobroff and increasing intake as palatability increased (see Table 2). All, Kissileff’s25study,subjectsfeltlesshungryandmoresatiated InternationalJournalofObesity Sensoryscienceandappetite LBSørensenetal 1156 after the palatable meal reflecting the larger amount Summary. The results from studies investigating the influ- eaten. Warwick et al10 found that after eating a palatable enceofpalatabilityonappetitesensationsinhumansgivean fixed preload, subjects felt less hungry and more full unclearpicture,asincreasingpalatabilityinonestudyresults compared to eating a less palatable preload. In Yeomans’18 in lower hunger and higher satiety ratings after consump- study from 1996, subjects felt no difference except for tion,butinotherstudiesresultsinhigherhungerandlower fullness,butthisreflectedtheamounteatenbythesubjects. fullness ratings or in higher hunger ratings and unaffected De Graaf et al19 observed lower hunger ratings after the fullnessratings,whereasothersshownodifferenceineffect higherpalatablepreload,reflectingthelargeramounteaten; of palatability. Studies also measuring the effect of palat- after eating the fixed preloads, there was no difference in ability on energy intake (the ‘within-meal’ effect) show effect of palatability. Yeomans and Symes11 observed no increasedintakeaspalatabilityincreases,nomatterwhatthe differences in ratings in spite of the higher intake of the effectofpalatabilityonthesubjectiveappetitesensationsis. palatable meal. Zandstra et al34 manipulated pleasantness The effect of palatability on postprandial appetite sensa- by varying the salt level and studied appetite over five tions after a test meal (the ‘after-meal’ effect) has been successive days, and at day 1 they observed higher ratings examinedinthreestudies,twoshowingastimulatingeffect for ‘faint with hunger’, ‘appetite for a snack’ and ‘appetite of palatability on subsequent appetite sensations and one for a meal’ for low-palatable bread meals than for medium study showing no difference. Two studies investigating the and high-palatable bread meals, and ratings for fullness effect of palatability on subsequent energy intake showed a were lower for the low-palatable bread meals, all ratings stimulating effect of palatability on energy intake and no reflecting the lower intake of low-palatable bread meal. differenceineffectofpalatability,respectively. During the 5-day period, this was changed and the investigators concluded that with repeated exposure, the Sensory-specificsatiety desiretoeatandintakeofalesspreferredfoodcouldincrease When a food is eaten, it drops in ratings of pleasantness overtime.34 relative to foods that have not been eaten and this InagreementwiththefindingsofHilletal8andMonneuse phenomenon is referred to as sensory-specific satiety.24 et al31, Rogers and Blundell36 observed that while there Althoughthetermsatietyisbeingused,appetitesensations were no differences in test meal intake and feelings of are not necessarily assessed. The standard procedure to test hunger and desire to eat after consumption of preloads sensory-specific satiety is to have moderately hungry of different palatability, during the subsequent 3h there subjects taste and rate the pleasantness of the sensory wasamorerapidrecoveryoffeelingsofhungeranddesireto propertiesofsmallportionsoffoods.Followingtheseinitial eat after the palatable preload compared both with the less ratings,oneofthefoodsisofferedasamealandsubjectseat palatablepreloadandnopreload.IncontrastDeGraafetal19 as much as they want. At various times following this test observed that neither hunger ratings nor food intake was meal, they taste and rate the foods tasted initially. Studies affected by the pleasantness of the test foods 2h after have shown that changes in the pleasantness of the foods consumption, and Warwick et al.10 observed that although occur immediately after consuming the test food and that there was a greater decrease in hunger ratings following the magnitude of these changes does not increase over thepalatablepreloads,thesensorypropertiesofthepreloads time.21,39,40Thisconfirmsthatthedevelopmentofsensory- did not influence the total caloric intake at lunch served specificsatietyisrelatedprimarilytothesensorystimulation 5hlater. accompanying ingestion as opposed to the postabsorptive All the above-mentioned studies examining the effect of effects of consuming these foods. The duration of sensory- palatability on appetite have been carried out in laboratory specific satiety is unknown; several studies show thatdrops environments, but we do not really know how much in liking persist more than 1h after consumption.39–41 understanding these studies provide about the influence of Sensory-specific satiety may also have an effect on the palatability on the normal food intake of humans in their decisiontoterminateameal.42 natural everyday environment. To investigate palatability Whatsensorypropertiesoffoodsareinvolvedinsensory- influences on eating behaviour, De Castro et al37 made 54 specific satiety? The best case is made for the gustatory French subjects maintain food intake diaries for four 7-day sensationsofsaltinessandsweetness.Eatingasavouryfood periods. The subjects recorded their intake along with tosatietyinvariablyleadstoadecreaseinthepleasantnessof palatabilityratingsofeachindividualitemeatenandalsoa that food and other savoury foods.20,23,43–45 The same global rating of palatability, which was an overall rating of phenomenon occurs when eating a sweet food to sati- the entire meal. The results showed that higher levels of ety.23,43–45 Other factors that could influence sensory- palatabilitywererelatedtolargermealsizes.Toextendthese specific satiety have been examined. To study whether findings, De Castro et al38 conducted a similar study with flavour intensity is involved in sensory-specific satiety, a 564 North Americans and this time the subjects recorded weak tea and a strong tea were compared and the test palatability only by global rating of the palatability of the showedanequivalentchangeinlikingforbothteas.46 entiremeal.Mealsthatwereratedhigherinpalatabilitywere Thepleasantnessofthesightofdifferentfoodsconsumed 44%largerthanmealslowinpalatability. tosatietydecreasedmorethanthepleasantnessofthesight InternationalJournalofObesity Sensoryscienceandappetite LBSørensenetal 1157 offoodsthathadnotbeeneaten.20Aftereatingsweetsofone approximatelyaslongasitwouldbeinthemouthduringa colour, the pleasantness of the taste of sweets eaten of that meal.48 particular colour declined more than that of the noneaten The role of different macronutrients in sensory-specific sweetsofothercolours,althoughthesweetsdifferedonlyin satiety has been examined and different results have been appearance.47Changesintheshapeofpasta(whichaffected reached. In several studies, no evidence of macronutrient boththeappearanceandthemouthfeelofthepasta)ledtoa specificitywasobtained,21,40,49–51butJohnsonandVickers52 specificdecreaseinthepleasantnessoftheshapeeaten.47In observedatrendforhigh-proteinfoodstodecreasemorein another study, texture-specific satiety was observed but not likingthanlow-proteinfoodsandVandewaterandVickers22 to the same extent as taste-specific satiety.23 Taste and foundthathigher-proteinfoods,wheneateninequalenergy olfactory sensory-specific satiety can be produced by chew- amounts, produced greater sensory-specific satiety than ing samples of a food or by smelling the food for lower-proteinfoods(seeTable3).Astudyoftheroleoffibre Table3 Sensory-specificsatietyFeffectofmacronutrients Study Subjects Preload Liking/pleasantness Hungerratings Typeoftestmeal Foodintake ratings Rollsetal21 10female Chicken,pasta, SSSafterallpreloads HungrierafterTurkish Self-selectionlunch Intakewaslessafter (1988) Withinsubjects creamcheeseon exceptchocolate.No Delight,creamcheese withfoodshighin chickenandpasta celery,TurkishDelight, differencesin andchocolatethan certainmacronutrients, chocolate pleasantnessofeaten afterpastaandchicken 2hdelay anduneatenfoodsof similarmacronutrient content Rollsetal49 16males Chocolatepudding NodifferencesinSSS Nodifferences Self-selectionmeal, Nodifferences (1989) 16females andstrawberry– 2hdelay }In2groups bananagelatine (info,noinfoa) desserts,highandlow calorie Johnsonand 13males None Thehigherprotein Theturkeymeal Cornwithbutter,cola NR Vickers52 25females mealsdroppedmorein suppressedappetite withliquidglucose (1992) Between likingthanthelower morethantheother polymer,cheese, subjects proteinmeals.Buttered meals butteredroll,turkey rollandcoladropped breast,potatochips, theleast sweets,canned peachesornomeal Johnsonand 6males 150 kcalor300kcal: Theeatenfood Pastaandchickenwere Self-selectionlunch, Lowerintakefollowing Vickers40 8females pastawithtomato appearedtoinfluence moresatiatingthanthe 90mindelay thechickenthanthe (1993) Withinsubjects sauce,blueberry- likingofuneatenfoods whippedcream whippedcream flavouredwhipped withsimilarflavours cream,chicken butnotsimilar macronutrients Vandewater 13/11males None SSSafterallpreloads. Thehigh-proteinmeals 1980kJ,sandwich, NR andVickers22 27/29females Thehigh-protein suppressedappetite highandlow-protein (1996) Withinsubjects yoghurtwasliked morethanthelow- 3240kJ,yoghurt,high betterthanthe proteinversions andlow-protein low-protein,yoghurt. HigherSSSwith high-proteinmeals comparedto low-proteinversions Milleretal50 Noinfob: Adlibregularpotato Noeffectofpotato NR Turkey,strawberry Nodifferencesin (2000) 24males chipsorpotatochips chiptypeonSSS yoghurt,cookieand intake(weight)of 20females withOlestraina carrot potatochipsonSSS Infob: 10-daysessioneach testdays,orbetween 27males thetwoinformation 24females conditions Withinsubjects SSS:sensory-specificsatiety;NR:notreported. aTheinformedgroupwasinformedofthenumberofcaloriesperservingandalsowhetherthefoodwashighorlowcalorie. bNoinfo/info ¼ theparticipantswerenotinformed/wereinformedaboutthefatandenergycontentofthechips. InternationalJournalofObesity Sensoryscienceandappetite LBSørensenetal 1158 in sensory-specific satiety showed no effect of a high-fibre diet could develop a long-term form of sensory-specific treatment vs a low-fibre treatment when muffins or satiety.54 soup were matched closely on sensory characteristics.53 In a study conducted by Kamphuis et al,51 fat-specific In two studies, Rolls et al41,49 observed that very low- satiety during dinner with oil high in linoleic acid was caloriefoodsproducedsensory-specificsatiety.Milleretal50 shown after a 2-week treatment, while there was no found that eating potato chips with high or low energy reductionintotalenergyintake. contentdidnotinfluencesensory-specificsatietydifferently. Summary. So far, flavour, texture and appearance-specific Whiletheseresultsindicatethatsensory-specificsatietydoes satietieshavebeenidentified,whereasstudiesontheroleof not depend on the energy content of the food, De Graaf et al45 and Johnson and Vickers40 observed that the extent macronutrients and the energy content of the food in sensory-specific satiety have given equivocal results. Differ- of sensory-specific satiety for an eaten food increased ent studies have shown that macronutrients and energy with increases in the amount of energy and/or food eaten content play a role in sensory-specific satiety or that (seeTable4). macronutrients and energy content are not a factor in In contrast to the studies described above where sensory- sensory-specificsatiety. specificsatietyhasbeeninvestigatedwithinasinglemeal,a long-term form of sensory-specific satiety has also been studied. Measures of the palatability of foods that the Varietyandfoodintake subjects had been eaten for many months was compared The sensory-specific satiety effect may have an important with measures of the palatability of other foods that the influenceontheamountoffoodeaten.Ifsatietyisspecificto subjects had not been eating; the subjects were refugees individualsensorypropertiesoffoods,thenintakeshouldbe from Ethiopia. The results indicated that monotony in the greater when a variety of foods are consumed together or Table4 Sensory-specificsatietyFeffectofenergy Study Subjects Preload Liking/pleasantness Hungerratings Typeoftestmeal Foodintake ratings Rollsetal41 24females TomatosouporJelly, NodifferencesinSSS Nodifferences Cheeseoncrackers, Nodifferencesinthe (1988) Withinsubjects highandlowcalorie 1hdelay weightsoffoodeaten. Highertotalenergy intakeinthehigh- calorie,soupcondition Rollsetal49 16males Chocolatepudding NodifferencesinSSS Nodifferences Self-selectionmeal, Nodifference (1989) 16females andstrawberry– 2hdelay In2groups bananagelatine (info,noinfoa) desserts,highandlow calorie DeGraaf Experiment1: Sweetandnonsweet NM Exp.1and2:sweet Exp.2:threebuffet Exp.1:NR etal45 28females foods250,350,450, preloadssuppressed meals:lunch,snack Exp.2:sweetpreloads (1993) Withinsubjects 550and650kcal appetiteforsomething anddinnerconsisting suppressedtheenergy Experiment2: Sweetandnonsweet sweetinadose– ofavarietyofsavoury intakefromsweetfood 29females foods250,500and responsemanner. andsweetfoods inadose–response Withinsubjects 750kcal Thesamecasefor manner,thesamecase nonsweetpreload fornonsweetpreload Johnson 6males 150or300kcal:pasta Highercaloriepreloads High-calorieloadswere Self-selectionlunch, Lowerintakefollowing and 8females withtomatosauce, droppedmoreinliking moresatiatingthan 90mindelay high-calorieloadsthan Vickers40 Withinsubjects blueberry-flavoured thanlower lowcalorie lowcalorie (1993) whippedcream, chicken Miller Noinfob: Adlibregularpotato Noeffectofpotato NR Turkey,strawberry Nodifferencesin etal50 24males chipsorpotatochips chiptypeonSSS yoghurt,cookieand intake(weight)of (2000) 20females withOlestraina carrot potatochipsonSSS Withinsubjects 10-daysessioneach testdays,orbetween Infob:27males thetwoinformation 24females conditions Withinsubjects SSS:sensory-specificsatiety. aTheinformedgroupwasinformedofthenumberofcaloriesperservingandalsowhetherthefoodwashighorlowcalorie. bNoinfo/info¼theparticipantswerenotinformed/wereinformedaboutthefatandenergycontentofthechips. InternationalJournalofObesity Sensoryscienceandappetite LBSørensenetal 1159 consecutively as opposed to when a single food is served. intakeeitheroverthecompletemealorinanyofthecourses Several studies have been carried out to investigate the afterthevarietywasintroduced.43Inastudyexaminingthe impact of meals varying in sensory properties such as impact of varying the colour of food, the presentation of a flavour,textureandappearance(seeTable5).Varyingflavour variety of colours of sweets did not affect food intake has shown diverse results. Changes in only the flavour of compared with consumption of the subjects’ favourite sandwich filling led to a 15% enhancement of food intake colour, although sensory-specific satiety was observed.47 In when all three flavours were presented successively com- the same study, changes in the shape of pasta led to a 14% pared with intake of the favourite flavour.47 On the other enhancementoffoodintakewhenthreeshapeswereoffered hand, offering subjects a variety of flavours of yoghurts of compared with intake of the subjects’ favourite shape.47 In the same colour and texture in succession did not enhance another study, subjects received hors d’oeuvres of one kind Table5 Effectsofavarietymealcomparedwithano-varietymeal Study Subjects Testmeal Hungerratings Foodintake Plineretal55(1980) 72malesF24normal Novariety(onetypehors Nodifferences Varietyincreasedintake weightnondieters d’oeuvres)orvariety(3kinds) Bellisleand 3males 5differentflavouredsandwiches NR Varietyincreasedintake LeMagnen28,29 7females (novariety),amixedmealinwhich (1980,1981) Withinsubjects all5flavourswereoffered(variety). Mealswereequicaloric Rollsetal43(1981) Experiment1: Experiment1:sandwicheswiththe NR Experiments1and2: 36females samefilling(plain)or4different Varietyincreasedintake Withinsubjects fillings(variety) Experiment2: Experiment2:yoghurtwith1 12females flavour(plain)or3different 12males flavours(variety) Withinsubjects Rollsetal47(1982) Inallexperiments: Exp.1:threeconditionswith NR Experiment1: 12males sweetsFmostpreferredcolour nodifference 12females (plain),adifferentcolouronein Experiments2and3: Withinsubjects eachcourseandequalmixofall varietyincreasedintake fourcolours Exp.2:tomatosaucewithpastain differentshapesalsointhree conditionsasexp.1 Exp.3:creamcheeseflavoured withsalt,lemonessenceand saccharin,orcurryspreadon bread.Uniformappearanceof spreadusingyellowcolour.Three conditionsasabove Rollsetal44(1984) 24males Twoconditions:mealwitha NR Varietyincreasedintake 24females differentfoodineachoffour Withinsubjects courses(varied)orthesamefood ineachcourse(plain) Berryetal56(1985) 65males 1flavouricecream,group;1 NR Varietyincreasedintake, 61females flavour,alone;3flavours,group;3 exceptformeneatingina Betweensubjects flavours,alone group Wisniewskietal57 40males 4conditions:pizza,cheeseburger, Nodifferences Varietyincreasedintake (1992) Betweensubjects pizza–pizza/cheeseburger– cheeseburger,pizza– cheeseburger/cheeseburger–pizza Porrinietal58(1997) Exp.1:13males Exp.1:uniformmealFomelette Exp.1:subjectsstoppedeating NR whensatiatedbutnotcompletely full Exp.2:10males Exp.2:variedmealFomelette Exp.2:subjectsstoppedeating Withinsubjects andbuffet-styleself-selectionmeal whensatiatedandcompletelyfull NR:notreported. InternationalJournalofObesity Sensoryscienceandappetite LBSørensenetal 1160 in a single condition and three kinds of hors d’oeuvres Differentstimuli distinctively different in appearance, flavour and texture in Toexaminewhetherthesensorypropertiesoffatalone,that the variety condition. The subjects consumed greater is ‘the fatty taste’, play a part in appetite regulation, the amounts in the variety condition than in the no-variety nonabsorbable fat substitute olestra was used to uncouple condition, yet they did not show greater decreases in the sensory properties of fat from fat absorption and reported hunger.55 In two studies, serving sandwiches with metabolism.Creamofbroccolisoupswasdevelopedinthree differentfillingsinducedanincreaseinmealsizecompared versions:fat-free,fat-free+olestraandhighfat.Subsequent toservingsandwicheswithjustonekindoffilling.28,29,43In energy intake was not affected by the sensory properties or anotherstudy,yoghurtsdifferinginflavour,appearanceand cognitivecuesassociatedwiththesoups.Thus,noevidence texturewereservedinthevarietyconditionandtherewasa indicated that fatty taste per se affects the regulation of 12.6% elevation of intake when the amount eaten in the energyintake.60Incontrast,Ceciletal61observedthatwhen variety condition was compared with the amount eaten of a high-fat and a high-carbohydrate soup were administered themostpreferredyoghurt.43Inastudy,wheresubjectswere intragastrically there was no difference in ratings of given a four-course lunch as either a varied meal with hungerandfullnessandfoodintakebetweenthetwosoups different food in each course, or as a plain meal with the whereaswhenthetwosoupswereingested,thehigh-fatsoup samefoodineachcourse,energyintakewaselevatedby60% suppressed hunger, induced fullness and tended to reduce in the varied meal primarily because of increased food energyintakeinasubsequenttestmealmorethanthehigh- consumption in the third and fourth courses.44 Another carbohydrate soup. The two soups were equienergetic and studyshowedthatsubjectsatemoreicecreamwhenoffered therewasnodifferenceinpleasantnessofthetwosoups. threekindsoficecreamdifferinginflavourandcolourthan Cornell et al62 demonstrated that individual’s food con- when offered only one kind of their own choosing.56 In a sumption could be affected by a period of pre-exposure to study,theinfluenceofintroducinganewfoodafterrepeated sensoryaspectsoffoodpriortoeating.Thecombinationofa presentationsofonefoodonfoodconsumptionwasstudied. brieftasteandsightofeitherpizzaoricecreampresenteda Pizzaorcheeseburgerwasserveddesignedtovaryinsensory fewminutespriortoeatingstimulatedsubjectstoeatbeyond properties such as flavour, texture and appearance. The the point of satiety. They also demonstrated that priming presentationofthenewfoodresultedinsignificantlygreater with a highly preferred food enhanced subsequent con- caloric consumption than another serving of the same sumption of that food relative to another equally preferred food (540 vs 180kJ) with no influence on hunger or but not primed food, even in satiated subjects. In a study fullness.57 In yet another study, the investigators observed investigating factors that promote the initiation of eating, that when a uniform meal was consumed, subjects stopped Marcelino et al63 observed that appetite for pizza increased eating when they felt satiated but not completely full, with hunger and after being visually exposed to the pizza, whereaswhenservedavariedmealtheydidnotstopeating both factors acting independently. Desire to eat pizza untiltheyfeltsatiatedandcompletelyfull.Theinvestigators depended on the visual quality of the pizza and also on concluded that in this condition, sensory characteristics of hunger.Incontrast,Fedoroffetal64foundthatpre-exposure foods played an important role in controlling food intake tothesmellofbakingpizzadidnotinfluencehungerratings and made the uniform meal more satiating thanthe varied andfoodconsumptioncomparedtonoexposureinsubjects one.58 thathadrefrainedfromeatingfor2hpriortotheexperiment. Examiningtheeffectofincreasingthevarietyofsensorily Inastudytestingtheeffectoftemperature,vegetablejuice distinct but nutritionally identical foods on appetite, food was used as a preload. Male subjects experienced a greater intakeandbodyweight,over7days,foodandenergyintake reduction in their desire to eat following the cold preloads increasedasthevarietyoffoodsincreased.Intakeamounted thanaftereitherthehotpreloadsornopreloadcondition.65 to1.57,1.76and1.97(cid:2) restingmetabolicrateonthelow-, Themenalsohadasignificantlylowerfoodintakefollowing medium-andhigh-varietytreatments,respectively,although the cold preload compared to the no-preload condition. therewasnodieteffectonsubjectivelyratedhunger.59 Temperatureofthepreloadsdidnotaffecteitherdesiretoeat orfoodintakeinfemalesubjects. Summary. Increasing the variety of sensorily distinct foods that are almost identical in composition can increase Sweetness and appetite. Besides the contribution of sweet- food and energy intake and in the short to medium term nesstopalatability,therehasbeenaninterestintheeffectof alter energy balance. In the majority of the reviewed sweettasteperseonappetite;however,theroleofsweettaste studies, subjective appetite sensations were either not in appetite regulation is controversial. One way to investi- reported or not monitored. Three studies have assessed gate the influence of sweet taste has been to uncouple subjectiveappetitesensationsandinallthreestudies,variety sweetnessandenergysoastoevaluatetherelativecontribu- increased intake, while there was no diet effect on hunger tionofeachfactortothecontrolofappetite.Bysubstitution ratings.55,57,58 ofanartificialsweetener forsugarinabeverageorfood,its Long-termeffectsofalteringthevarietyoffoodsonenergy energyvalueisreducedwhileitssweetnessismaintainedand balanceareyettobedetermined. thentheeffectofthesweetenedbeverageorfoodonappetite InternationalJournalofObesity Sensoryscienceandappetite LBSørensenetal 1161 canbecomparedwiththeeffectofunsweetenedbeverageor that after administration of a benzodiazepine-receptor food(seeTable6).BlundellandHill66reportedanincreasein agonist, the number of eating occasions increased without motivationtoeatanddecreasedratingsoffullnessfollowing alteringmealsize.Palatability,however,wasnotassessed. the ingestion of an aspartame solution compared to inges- Studies investigating the hypothesis that palatability and tion of water, which is consistent with the hypothesis that sweet taste have an effect on cephalic phase insulin release sweet taste can stimulate hunger. Support for this view has anddiet-inducedthermogenesishaveshowndifferentresults. come from other short-term studies showing a stimulating In some studies, the cephalic phase plasma insulin levels effect of sweet taste on hunger ratings when the sweetener measured were lower when subjects were consuming an wasprovidedintheformofaqueoussolutions,chewinggum unpalatablemealthanwhenconsumingpalatablefood,92–96 orbeverages9,67–69andbothhungerratingsandfoodintake whileinastudyusingshamfeeding,nosignificantdifference when using fruit yoghurt as avehicle.70In contrast, several inthemagnitudeofcephalicphaseinsulinreleasewasfound studies showed no effect of sweet taste on either hunger between palatable and unpalatable food.97 Studies on the ratings or food intake when the sweetener was provided in effect of sweet taste per se on cephalic phase insulin release theformofgelatine,cornflakesorfromageblanc71–74oras failedtoshowinsulinsecretionafteroralstimulationofsweet aspartame-orsaccharin-sweeteneddrinks.75–79 taste.98–100Studiesonthe effect of diet-induced thermogen- Other studies have contributed to elucidate the role of esisofpalatableandunpalatablefoodshaveshowndifferent sweettasteonappetite.BralaandHagan80showedthatwhen results. In some studies, diet-induced thermogenesis was subjects’ ability to taste sweetness was attenuated by significantlyelevatedafterconsumptionofapalatablemeal gymnemic acid, they ate less food after a sweet-tasting in comparison with an unpalatable meal,92,99,101,102 while preloadthandidsubjectswhosetasteperceptionwasintact. othersshowednodifferenceinthermogenesis.96,103Investi- Thesefindingssuggestthatasweetoralstimulationinitiates gation of the effect of sweet taste on diet-induced thermo- reflexesthatincreaseappetite.Rollsetal81havestudiedthe genesisshowedthatsweettastehasnoeffect.104 effect of eating a sweet food either as a first course or as a It has been speculated that sensory-specific satiety is thirdcourseonfoodintakeduringathree-courselunch.The caused by sensory adaptation, that is, the intensity of a secondcourseremainedthesameinthetwoconditionsand food’s flavour diminishes when there is a sensory-specific thetwocoursesthatweretobeinterchangedwereofsimilar decreaseinthepleasantnessofthefood.Thisdoesnotseem energydensitiesandratedpalatability,onecourseconsisted to be the case, as studies have shown that changes in the of a chocolate bar and the other was cheese on crackers. pleasantnessofafooddonotappeartoreflectchangesinthe There was no indication that eating candy as a first course perceivedintensityofthefoods.20,48Insteadofbeingrelated affectedappetiteandfoodintakedifferentlyfromeatingthe to changes in the sensory processing of responses to foods, same amount of a cheese on crackers. In two experiments, sensory-specific satiety is more likely related to brain areas De Graaf et al45 studied the hypothesis that if sweet taste controllingmotivationandtherewardvalueoffoods.105–108 sensation increases appetite then sweet carbohydrates must belesssatiatingthannonsweetcarbohydrates.Theresultsof thefirstexperimentwereconsistentwiththehypothesis,but Suggestions for future work thesecondexperimentdidnotshowanydifferencebetween Results reviewed in this paper suggest that increased sweetandnonsweetcarbohydrates. palatability increases food intake in the short term. The contradictory results found when assessing subjective appe- titesensationsmaybearesultofdifferentstudydesigns,but Summary. The role of sweet taste per se in appetite itisworthstudyingwhetherthesediverseresultsinfactare regulationisnotfullyclarified.Althoughsomestudiesshow due to different methodologies or whether there is a a stimulating effect, most studies show no effect of sweet discrepancy between subjective appetite sensations and tasteonappetiteandfoodintake. actualfoodintakewheneatingapalatablemeal. Sensory-specific satiety may influence the amount of food Mechanisms eaten,asincreasingthevarietyoffoodsincreasesfoodintake. What physiological mechanisms are behind the possible Thus,itmaybethatvarietyenhancesthepalatabilityofameal effect of palatability on appetite and food intake? Study and people therefore eat more. Here, as in the palatability results imply that endogenous opioid peptides are involved studies, some interesting results appeared when subjective inprocessesunderlyingthedeterminationofpalatability.In appetite sensations were assessed: in three studies, variety several studies using opioid antagonists, intake is reduced, increasedfoodintakewithoutinfluencingsubjectivehunger butmainlyforthefoodsthatareratedashighlypalatable.82–86 ratings.Thissupportstheideathattheamountofpalatable Hetherington et al,87 however, failed to find an effect of food eaten may not be recognised and therefore subjective opioidantagonismonpleasantnessratingsandintake.Accord- appetiteratingsdonotreflecttheamountoffoodeaten. ing to several animal studies, benzodiazepine may enhance Long-termstudiesoftheconsequencesoffoodpalatability food palatability and thereby increase meal size.88–90 The andvarietyonfoodintakeandenergybalancecarriedoutin onlystudyonhumans,performedbyHaneyetal,91showed everydaynaturalenvironmentsareneeded. InternationalJournalofObesity

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