There is currently great interest in developing ecological models for investigating human evolution. Yet little attention has been given to energetics, one of the cornerstones of modern ecosystem ecology. This paper examines the ecological correlates of variation in metabolic requirements among extant primate species, and uses this information to draw inferences about the changes in energy demands over the course of human evolution. Data on body size, resting metabolism, and activity budgets for selected anthropoid species and human hunter-gatherers are used to estimate total energy expenditure (TEE). Analyses indicate that relative energy expenditure levels and day ranges are positively correlated with diet quality; that is, more active species tend to consume more energy-rich diets. Human foragers fall at the positive extremes for modern primates in having high expenditure levels, large ranges, and very high quality diets. During hominid evolution, it appears that TEE increased substantially with the emergence of Homo erectus. This increase is partly attributable to larger body size as well as likely increases in day range and activity level. Assuming similar activity budgets for all early hominid species, estimated TEE for H. erectus is 40-45% greater than for the australopithecines. If, however, it is assumed that the evolution of early Homo was also associated with a shift to a more "human-like" foraging strategy, estimated expenditure levels for H. erectus are 80-85% greater than in the australopithecines. Changing patterns of resource distribution associated with the expansion of African savannas between 2.5 and 1.5 mya may been the impetus for a shift in foraging behavior among early members of the genus Homo. Such ecological changes likely would have made animal foods a more attractive resource. Moreover, greater use of animal foods and the resulting higher quality diet would have been important for supporting the larger day ranges and greater energy requirements that appear to have been associated with the evolution of a human-like hunting and gathering strategy.

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Human dietary patterns and metabolic requirements are compared to those of nonhuman primate species in order to gain insights into the evolution of our nutritional needs. In general, primate diet quality (i.e., caloric and nutrient density) is inversely related to body size and total resting metabolic requirements (RMR). Humans, however, consume a diet of much higher quality than is expected for our size and metabolic needs. This energy-rich diet appears to reflect an adaptation to the high metabolic cost of our large brain. Among primates, the relative proportion of resting metabolic energy used for brain metabolism is positively correlated with relative diet quality. Humans represent the positive extreme, having both a very high quality diet and a large brain that accounts for 20-25% of resting metabolism. Evidence from the hominid fossil record implies that major changes in diet and relative brain metabolism occurred with the emergence of the genus Homo. © 1994 Wiley-Liss, Inc.

A bioenergetics model is developed to examine changes in metabolic requirements over the course of human evolution. Data on (1) body size and resting metabolism, (2) brain size and metabolism, (3) activity budgets, and (4) foraging patterns for humans and other anthropoids are used to evaluate ecological correlates of variation in diet and energy expenditure. Analyses of variation in these extant species provide a framework for estimating (1) resting metabolic requirements, (2) brain metabolic needs, and (3) total energy requirements in fossil hominids. Anthropoid primates spend about 8% of resting metabolism to maintain their brains, a significantly larger proportion than in other mammals (3-4%), but still significantly less than 20-25% in humans. Total energy expenditure among anthropoids is positively correlated with day range and dietary quality. Human foragers fit this pattern, having high levels of energy expenditure, large foraging ranges, and a high quality diet. Within the fossil record, it appears that both total energy expenditure (TEE) and energy required by the brain increased substantially with the emergence of Homo erectus. For H. erectus, the percentage of resting metabolism used by the brain falls beyond the nonhuman primate range and approaches the modern human range. Additionally, TEE is 35-55% greater than in the australopithecines. The high total metabolic needs and the large proportion of energy required by the brain imply that important dietary changes occurred with H. erectus. These metabolic and dietary changes are linked to (1) the emergence of hunting and gathering, (2) the evolution of the human pattern of prolonged development, and (3) the coexistence and competition with the robust australopithecines.

The evolution of large human brain size has had important implications for the nutritional biology of our species. Large brains are energetically expensive, and humans expend a larger proportion of their energy budget on brain metabolism than other primates. The high costs of large human brains are supported, in part, by our energy- and nutrient-rich diets. Among primates, relative brain size is positively correlated with dietary quality, and humans fall at the positive end of this relationship. Consistent with an adaptation to a high-quality diet, humans have relatively small gastrointestinal tracts. In addition, humans are relatively "undermuscled" and "over fat" compared with other primates, features that help to offset the high energy demands of our brains. Paleontological evidence indicates that rapid brain evolution occurred with the emergence of Homo erectus 1.8 million years ago and was associated with important changes in diet, body size, and foraging behavior.