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Am J Clin Nutr 2005 Cordain 341 54

Am J Clin Nutr 2005 Cordain 341 54

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Commentary
Origins and evolution of the Western diet: health implications for the21st century
1,2
 Loren Cordain, S Boyd Eaton, Anthony Sebastian, Neil Mann, Staffan Lindeberg, Bruce A Watkins, James H O’Keefe,and Janette Brand-Mille
ABSTRACT
There is growing awareness that the profound changes in the envi-ronment (eg, in diet and other lifestyle conditions) that began withtheintroductionofagricultureandanimalhusbandry
Ȃ
10 000yagooccurred too recently on an evolutionary time scale for the humangenome to adjust. In conjunction with this discordance between ourancient,geneticallydeterminedbiologyandthenutritional,cultural,andactivitypatternsofcontemporaryWesternpopulations,manyof the so-called diseases of civilization have emerged. In particular,food staples and food-processing procedures introduced during theNeolithic and Industrial Periods have fundamentally altered 7 cru-cial nutritional characteristics of ancestral hominin diets:
1
) glyce-mic load,
2
) fatty acid composition, 3) macronutrient composition,
4
)micronutrientdensity,
5
)acid-basebalance,6)sodium-potassiumratio, and
) fiber content. The evolutionary collision of our ancientgenomewiththenutritionalqualitiesofrecentlyintroducedfoodsmayunderliemanyofthechronicdiseasesofWesterncivilization.
Am J Clin Nutr 
2005;81:341–54.
KEY WORDS
Westernized diets, chronic disease, processedfoods, genetic discordance, hunter-gatherers, human evolution
EVOLUTIONARY DISCORDANCE
Evolution acting through natural selection represents an on-goinginteractionbetweenaspecies’genomeanditsenvironmentover the course of multiple generations. Genetic traits may bepositively or negatively selected relative to their concordance ordiscordance with environmental selective pressures (1). Whenthe environment remains relatively constant, stabilizing selec-tion tends to maintain genetic traits that represent the optimalaverage for a population (2). When environmental conditionspermanently change, evolutionary discordance arises between aspecies’genomeanditsenvironment,andstabilizingselectionisreplacedbydirectionalselection,movingtheaveragepopulationgenome to a new set point (1, 2). Initially, when permanentenvironmental changes occur in a population, individuals bear-ing the previous average status quo genome experience evolu-tionary discordance (2, 3). In the affected genotype, this evolu-tionary discordance manifests itself phenotypically as disease,increased morbidity and mortality, and reduced reproductivesuccess (1–3).Similar to all species, contemporary humans are geneticallyadapted to the environment of their ancestors—that is, to theenvironment that their ancestors survived in and that conse-quently conditioned their genetic makeup (1–3). There is grow-ing awareness that the profound environmental changes (eg, indiet and other lifestyle conditions) that began with the introduc-tion of agriculture and animal husbandry
Ȃ
10 000 y ago oc-curred too recently on an evolutionary time scale for the humangenome to adapt (2–5). In conjunction with this discordancebetween our ancient, genetically determined biology and thenutritional,cultural,andactivitypatternsincontemporaryWest-ern populations, many of the so-called diseases of civilizationhave emerged (2–12).
CHRONIC DISEASE INCIDENCE
In the United States, chronic illnesses and health problemseither wholly or partially attributable to diet represent by far themost serious threat to public health. Sixty-five percent of adultsaged
ͧ
20 y in the United States are either overweight or obese(13), and the estimated number of deaths ascribable to obesity is280 184peryear(14).Morethan64millionAmericanshaveoneormoretypesofcardiovasculardisease(CVD),whichrepresentstheleadingcauseofmortality(38.5%ofalldeaths)intheUnitedStates(15).FiftymillionAmericansarehypertensive;11millionhave type 2 diabetes, and 37 million adults maintain high-risk total cholesterol concentrations (
240 mg/dL) (15). In post-menopausal women aged
ͧ
50 y, 7.2% have osteoporosis and39.6% have osteopenia (16). Osteoporotic hip fractures are as-sociated with a 20% excess mortality in the year after fracture
1
From the Department of Health and Exercise Science, Colorado StateUniversity, Fort Collins (LC); the Departments of Radiology and Anthro-pology, Emory University, Atlanta (SBE); the Department of Medicine andUCSF/Moffitt General Clinical Research Center, University of California,San Francisco (AS); the Department of Food Science, RMIT University,Melbourne, Australia (NM); the Department of Medicine, Lund University,Sweden (SL); the Department of Food Science, Lipid Chemistry and Mo-lecular Biology Laboratory, Purdue University, West Lafayette, IN (BAW);the Mid America Heart Institute, Cardiovascular Consultants, Kansas City,MO (JHO); and the Human Nutrition Unit, Department of Biochemistry,University of Sydney, Australia (JB-M).
2
Address reprint requests to L Cordain, Department of Health and Exer-cise Science, Colorado State University, Fort Collins, CO 80523. E-mail:cordain@cahs.colostate.edu.Received June 17, 2004.Accepted for publication August 24, 2004.
341
 Am J Clin Nutr 
2005;81:341–54. Printed in USA. © 2005 American Society for Clinical Nutrition
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(17). Cancer is the second leading cause of death (25% of alldeaths)intheUnitedStates,andanestimatedone-thirdofallcancerdeaths are due to nutritional factors, including obesity (18).
HOMININ DIETARY CHARACTERISTICS
In the 5–7 million-year period since the evolutionary emer-gence of hominins (bipedal primates within the taxonomic tribehominini; note that the newer term
hominin
supplants the previ-ous term,
hominid 
)
ͧ
20 species may have existed (
Figure 1
)(19). Similar to historically studied hunter-gatherers (20, 21),there would have been no single universal diet consumed by allextinct hominin species. Rather, diets would have varied bygeographic locale, climate, and specific ecologic niche. How-ever, there are universal characteristics of preagricultural homi-nindietsthatareusefulinunderstandinghowthecurrentWesterndiet may predispose modern populations to chronic disease. In-creasingly,clinicaltrialsandinterventionsthatusedietarytreat-ments with nutritional characteristics similar to those found inpreindustrial and preagricultural diets have confirmed the bene-ficial health consequences predicted by the template of evolu-tionary discordance theory.
NUTRITIONAL CHARACTERISTICS OF PRE- ANDPOSTAGRICULTURAL DIETS
Before the development of agriculture and animal husbandryhominin dietary choices would have been necessarily limited tominimally processed, wild plant and animal foods. With theinitial domestication of plants and animals, the original nutrientcharacteristics of these formerly wild foods changed, subtly atfirst but more rapidly with advancing technology after the In-dustrialRevolution.Furthermore,withtheadventofagriculture,novel foods were introduced as staples for which the hominingenome had little evolutionary experience. More importantly,food-processing procedures were developed, particularly fol-lowingtheIndustrialRevolution,whichallowedforquantitativeand qualitative food and nutrient combinations that had not pre-viously been encountered over the course of hominin evolution.Incontrastingpre-andpostagriculturaldiets,itisimportanttoconsider not only the nutrient qualities and types of foods thatlikely would have been consumed by preagricultural homininsbuttoalsorecognizethetypesoffoodsandtheirnutrientqualitiesthat could not have been regularly consumed before the devel-opment of agriculture, industrialization, and advanced technol-ogy. Food types that would have generally been unavailable topreagriculturalhomininsarelistedin
Table1
(22–24).Althoughdairy products, cereals, refined sugars, refined vegetable oils,andalcoholmakeup72.1%ofthetotaldailyenergyconsumedbyall people in the United States, these types of foods would havecontributed little or none of the energy in the typical preagricul-tural hominin diet (20). Additionally, mixtures of foods listed inTable 1 make up the ubiquitous processed foods (eg, cookies,cake, bakery foods, breakfast cereals, bagels, rolls, muffins,crackers, chips, snack foods, pizza, soft drinks, candy, ice cream,condiments,andsaladdressings)thatdominatethetypicalUSdiet.
Dairy foods
Hominins, like all mammals, would have consumed the milk of their own species during the suckling period. However, afterweaning, the consumption of milk and milk products of othermammalswouldhavebeennearlyimpossiblebeforethedomes-tication of livestock because of the inherent difficulties in cap-turingandmilkingwildmammals.Althoughsheepweredomes-ticatedby
Ȃ
11 000beforepresent(BP)(25)andgoatsandcowsby
Ȃ
10 000 BP (26, 27), early direct chemical evidence fordairying dates to 6100 to 5500 BP from residues of dairy fatsfound on pottery in Britain (28). Taken together, these data in-dicate that dairy foods, on an evolutionary time scale (Figure 1),are relative newcomers to the hominin diet.
FIGURE 1.
The hominin fossil record. Species are indicated with the dates of the earliest and latest fossil record. Adapted from Wood (19).
342
CORDAIN ET AL
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Cereals
Because wild cereal grains are usually small, difficult to har-vest,andminimallydigestiblewithoutprocessing(grinding)andcooking, the appearance of stone processing tools in the fossilrecord represents a reliable indication of when and where cul-tures systematically began to include cereal grains in their diet(7).Groundstonemortars,bowls,andcupholesfirstappearedinthe Upper Paleolithic (from 40 000 y ago to 12 000 y ago) (29),whereas the regular exploitation of cereal grains by any world-wide hunter-gatherer group arose with the emergence of theNatufian culture in the Levant
Ȃ
13 000 BP (30). DomesticationofemmerandeinkornwheatbythedescendantsoftheNatufiansheralded the beginnings of early agriculture and occurred by10–11 000BPfromstrainsofwildwheatlocalizedtosoutheast-ern Turkey (31). During the ensuing Holocene (10 000 y agountil the present), cereal grains were rarely consumed as yearround staples by most worldwide hunter-gatherers (32, 33), ex-cept by certain groups living in arid and marginal environments(32, 34). Hence, as was the case with dairy foods, before theEpi-Paleolithic (10 000–11 000 y ago) and Neolithic (10 000 to5500 y ago) periods, there was little or no previous evolutionaryexperience for cereal grain consumption throughout homininevolution.In Table 1, it is shown that 85.3% of the cereals consumed inthe current US diet are highly processed refined grains. Preced-ingtheIndustrialRevolution,allcerealsweregroundwiththeuseof stone milling tools, and unless the flour was sieved, it con-tainedtheentirecontentsofthecerealgrain,includingthegerm,bran, and endosperm (35). With the invention of mechanizedsteel roller mills and automated sifting devices in the latter partof the 19th century (35), the nutritional characteristics of milledgrain changed significantly because the germ and bran wereremoved in the milling process, leaving flour comprised mainlyof endosperm of uniformly small particulate size (35, 36). Ac-cordingly, the widespread consumption of highly refined grainflours of uniformly small particulate size represents a recentsecular phenomenon dating to the past 150–200 y (35).
Refined sugars
ThepercapitaconsumptionofallrefinedsugarsintheUnitedStates in 2000 was 69.1 kg, whereas in 1970 it was 55.5 kg (24).ThisseculartrendforincreasedsugarconsumptionintheUnitedStatesinthepast30yreflectsamuchlargerworldwidetrendthathas occurred in Western nations since the beginning of the In-dustrial Revolution some 200 y ago (37). The per capita refinedsucrose consumption in England steadily rose from 6.8 kg in1815to54.5kgin1970(38),asshownin
Figure2
.Similartrendsin refined sucrose consumption have been reported during theIndustrial Era for the Netherlands, Sweden, Norway, Denmark,and the United States (39).The first evidence of crystalline sucrose production appearsabout 500 BC in northern India (37). Before this time, honeywould have represented one of the few concentrated sugars towhich hominins would have had access. Although honey likelywas a favored food by all hominin species, seasonal availabilitywould have restricted regular access. Studies of contemporaryhunter-gatherers show that gathered honey represented a rela-tivelyminordietarycomponentoverthecourseofayear,despitehigh intakes in some groups during short periods of availability.In the Anbarra Aborigines of northern Australia, average honeyconsumptionoverfour1-moperiods,chosentoberepresentativeof the various seasons, was 2 kg per person per year (40). In theAche Indians of Paraguay, honey represented 3.0% of the aver-age total daily energy intake over 1580 consumer days (41).Consequently,currentpopulation-wideintakesofrefinedsugarsin Westernized societies represent quantities with no precedentduring hominin evolution.Inthepast30y,qualitativefeaturesofrefinedsugarconsump-tion have changed concurrently with the quantitative changes.With the advent of chromatographic fructose enrichment tech-nology in the late 1970s, it became economically feasible tomanufacture high-fructose corn syrup (HFCS) in mass quantity
TABLE 1
Food and food types found in Western diets generally unavailable topreagricultural hominins
1
Food or food group ValueDairy products
% of energy
2
Whole milk 1.6Low-fat milk 2.1Cheese 3.2Butter 1.1Other 2.6Total 10.6Cereal grainsWhole grains 3.5Refined grains 20.4Total 23.9Refined sugarsSucrose 8.0High-fructose corn syrup 7.8Glucose 2.6Syrups 0.1Other 0.1Total 18.6Refined vegetable oilsSalad, cooking oils 8.8Shortening 6.6Margarine 2.2Total 17.6Alcohol 1.4Total energy 72.1Added salt, as sodium chloride 9.6
31
Data adapted from references 22–24.
2
In the US diet.
3
Salt from processed foods, table salt use, and cooking; in g/d.
FIGURE 2.
Per capita consumption of sucrose in England from 1815 to1970.WWI,WorldWarI;WWII,WorldWarII.AdaptedfromCleave(38).
WESTERN DIETARY ORIGINS
343
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