American Journal of Clinical Nutrition, Vol. 71, No. 3, 682-692, March 2000
© 2000 American Society for Clinical Nutrition

--------------------------------------------------------------------------------

Special Article

Plant-animal subsistence ratios and macronutrient energy estimations in worldwide hunter-gatherer diets1,2

Loren Cordain, Janette Brand Miller, S Boyd Eaton, Neil Mann, Susanne HA Holt and John D Speth

1 From the Department of Health and Exercise Science, Colorado State University, Fort Collins; the Human Nutrition Unit, Department of Biochemistry, University of Sydney, New South Wales, Australia; the Departments of Radiology and Anthropology, Emory University, Atlanta; the Department of Food Science, Royal Melbourne Institute of Technology University, Melbourne, Australia; and the Museum of Anthropology, University of Michigan, Ann Arbor.

Both anthropologists and nutritionists have long recognized that the diets of modern-day hunter-gatherers may represent a reference standard for modern human nutrition and a model for defense against certain diseases of affluence. Because the hunter-gatherer way of life is now probably extinct in its purely un-Westernized form, nutritionists and anthropologists must rely on indirect procedures to reconstruct the traditional diet of preagricultural humans. In this analysis, we incorporate the most recent ethnographic compilation of plant-to-animal economic subsistence patterns of hunter-gatherers to estimate likely dietary macronutrient intakes (% of energy) for environmentally diverse hunter-gatherer populations. Furthermore, we show how differences in the percentage of body fat in prey animals would alter protein intakes in hunter-gatherers and how a maximal protein ceiling influences the selection of other macronutrients. Our analysis showed that whenever and wherever it was ecologically possible, hunter-gatherers consumed high amounts (45–65% of energy) of animal food. Most (73%) of the worldwide hunter-gatherer societies derived >50% (56–65% of energy) of their subsistence from animal foods, whereas only 14% of these societies derived >50% (56–65% of energy) of their subsistence from gathered plant foods. This high reliance on animal-based foods coupled with the relatively low carbohydrate content of wild plant foods produces universally characteristic macronutrient consumption ratios in which protein is elevated (19–35% of energy) at the expense of carbohydrates (22–40% of energy).

-------------------------------
: Eur J Clin Nutr 2002 Mar;56 Suppl 1:S42-52

The paradoxical nature of hunter-gatherer diets: meat-based, yet non-atherogenic.

Cordain L, Eaton SB, Miller JB, Mann N, Hill K.

Department of Health and Exercise Science, Colorado State University, Fort Collins, Colorado, USA. cordain~cahs.colostate.edu

OBJECTIVE: Field studies of twentieth century hunter-gathers (HG) showed them to be generally free of the signs and symptoms of cardiovascular disease (CVD). Consequently, the characterization of HG diets may have important implications in designing therapeutic diets that reduce the risk for CVD in Westernized societies. Based upon limited ethnographic data (n=58 HG societies) and a single quantitative dietary study, it has been commonly inferred that gathered plant foods provided the dominant energy source in HG diets. METHOD AND RESULTS: In this review we have analyzed the 13 known quantitative dietary studies of HG and demonstrate that animal food actually provided the dominant (65%) energy source, while gathered plant foods comprised the remainder (35%). This data is consistent with a more recent, comprehensive review of the entire ethnographic data (n=229 HG societies) that showed the mean subsistence dependence upon gathered plant foods was 32%, whereas it was 68% for animal foods. Other evidence, including isotopic analyses of Paleolithic hominid collagen tissue, reductions in hominid gut size, low activity levels of certain enzymes, and optimal foraging data all point toward a long history of meat-based diets in our species. Because increasing meat consumption in Western diets is frequently associated with increased risk for CVD mortality, it is seemingly paradoxical that HG societies, who consume the majority of their energy from animal food, have been shown to be relatively free of the signs and symptoms of CVD.
CONCLUSION: The high reliance upon animal-based foods would not have necessarily elicited unfavorable blood lipid profiles because of the hypolipidemic effects of high dietary protein (19-35% energy) and the relatively low level of dietary carbohydrate (22-40% energy). Although fat intake (28-58% energy) would have been similar to or higher than that found in Western diets, it is likely that important qualitative differences in fat intake, including relatively high levels of MUFA and PUFA and a lower omega-6/omega-3 fatty acid ratio, would have served to inhibit the development of CVD. Other dietary characteristics including high intakes of antioxidants, fiber, vitamins and phytochemicals along with a low salt intake may have operated synergistically with lifestyle characteristics (more exercise, less stress and no smoking) to further deter the development of CVD.

---------------------------------

American Journal of Clinical Nutrition, Vol. 72, No. 6, 1589-1590, December 2000
© 2000 American Society for Clinical Nutrition

--------------------------------------------------------------------------------

Letters to the Editor

Macronutrient estimations in hunter-gatherer diets
Loren Cordain, Janette Brand Miller, S Boyd Eaton and Neil Mann
Department of Health and Exercise Science Colorado State University Fort Collins, CO 80523 E-mail: cordain~cahs.colostate.edu
Department of Biochemistry University of Sydney Sydney, NSW 2006 Australia
Departments of Radiology and Anthropology Emory University Atlanta, GA 30322
Department of Food Science Royal Melbourne Institute of Technology University GPO Box 2476V Melbourne, VIC 3001 Australia

Dear Sir:

We disagree with the editorial (1) that accompanied our recent article on hunter-gatherer plant-animal subsistence ratios (2). Milton appears to have misinterpreted our findings as well as Lee's (3) original analysis of the Ethnographic Atlas (4).

Within the nutritional community, it is common knowledge that the quantitative and qualitative lipid composition of domesticated meats is vastly different from that found in wild game. Game meat contains lower proportions of fat, especially saturated fat, than does meat from grain-fed domesticated animals, even on a whole-carcass basis (5). Nowhere in our article did we recommend that people should eat high-fat, domesticated livestock. Our take-home messages were that hunter-gatherer diets were higher in protein and lower in carbohydrate than are current Western diets or dietary guidelines and that this macronutrient balance may provide insight into potentially therapeutic diets. If any implication were to be inferred, it would be that dietary fat should emulate fat sources found in game meat and organs (high in n-3 fats, low in n-6 fats, and high in monounsaturated fats).

Milton's editorial repeated the same error that has occurred continually in the anthropologic community since Lee published his work 32 y ago (3). Lee did not report the total food intakes derived from animal sources because he did not sum hunted and fished animal foods. This is one of the reasons our reanalysis of the Ethnographic Atlas is original and noteworthy. Although we did not report it in our article, we analyzed Lee's sample of 58 hunter-gatherer societies as a subset and obtained results almost identical to those of our analysis of the entire sample (n = 229). The dependence on hunted and fished foods for subsistence was 86–100% (modal value) and 66–75% (median value). Milton's statement that "emphasis on hunting occurred only in the highest latitudes" is also inaccurate because our analysis of Lee's data showed that there is no correlation (Spearman's rho = 0.01) between dependence on hunting and latitude; on the contrary: as intakes of plant food decrease with increasing latitude, intakes of fished food increase and of hunted animal food stay constant—the same conclusion we reached with our original analysis. The editorial deemphasizes the importance of animal foods in hunter-gatherer diets by citing 2 extreme and nonrepresentative societies, the !Kung and the Hazda, both of which have been shown by the Ethnographic Atlas and modern quantitative studies to maintain high plant-animal subsistence ratios (67:33 and 56:44, respectively). Of the 229 hunter-gatherer societies listed in the Ethnographic Atlas, only 1 other society maintains a plant-animal subsistence ratio as high as that of the !Kung and only 13% maintain a ratio as high or higher than that of the Hazda. A compilation of the few available quantitative dietary studies in hunter-gatherers showed a plant-animal subsistence ratio of 41:59 (6), which is similar to the aggregate value (45:55) we reported in our article.

Increases in low-fat dietary protein at the expense of carbohydrate may have therapeutic effects. Wolfe and Piche (7) showed that the replacement of dietary carbohydrate with low-fat, high-protein animal foods improved blood lipids (LDL, VLDL, total cholesterol, triacylglycerol, and the ratio of total to HDL cholesterol). Furthermore, increased dietary protein may reduce the risk of coronary heart disease (8) and reduce serum homocysteine concentrations (9) while facilitating weight loss (10) and improving insulin metabolism (11).

Again, we do not recommend increases in intakes of domesticated animal fat, only of lean protein from lean animals, preferably protein that may also contain significant amounts of n-3 and monounsaturated fat such as that found in game meat. Consumption of low-fat dietary protein at the expense of carbohydrate is the nutritional pattern that is consistent with our species' evolutionary history and represents a viable dietary option for improving health and well-being in modern people. Further research is needed before this dietary pattern can be recommended without reservation, particularly in subjects with preexisting kidney disease.

REFERENCES


Milton K. Hunter-gatherer diets—a different perspective. Am J Clin Nutr 2000;71:665–7 (editorial).[Full Text]
Cordain L, Brand Miller J, Eaton SB, Mann N, Holt SHA, Speth JD. Plant-animal subsistence ratios and macronutrient energy estimations in worldwide hunter-gatherer diets. Am J Clin Nutr 2000; 71:682–92.[Abstract/Full Text]
Lee RB. What hunters do for a living, or how to make out on scarce resources. In: Lee RB, DeVore I, eds. Man the hunter. Chicago: Aldine, 1968:30–48.
Murdock GP. Ethnographic atlas: a summary. Ethnology 1967;6: 109–236.
Eaton SB. Humans, lipids and evolution. Lipids 1992;27:814–20.[Medline]
Leonard WR, Robertson ML. Evolutionary perspectives on human nutrition: the influence of brain and body size on diet and metabolism. Am J Hum Biol 1994;6:77–88.
Wolfe BMJ, Piche LA. Replacement of carbohydrate by protein in a conventional-fat diet reduces cholesterol and triglyceride concentrations in healthy normolipidemic subjects. Clin Invest Med 1999; 22:140–8.[Medline]
Hu FB, Stampfer MJ, Manson JE, et al. Dietary protein and risk of ischemic heart disease in women. Am J Clin Nutr 1999;70:221–7.[Abstract/Full Text]
Mann NJ, Li D, Sinclair AJ, et al. The effect of diet on plasma homocysteine concentrations in healthy male subjects. Eur J Clin Nutr 1999;53:895–9.[Medline]
Skov AR, Toubro S, Ronn B, Holm L, Astrup A. Randomized trial on protein vs carbohydrate in ad libitum fat reduced diet for the treatment of obesity. Int J Obes Relat Metab Disord 1999;23:528–36.[Medline]
O'Dea K, Traianedes K, Ireland P, et al. The effects of diet differing in fat, carbohydrate, and fiber on carbohydrate and lipid metabolism in type II diabetes. J Am Diet Assoc 1989;89:1076–86.[Medline]