The idea that aging should be linked to energy expenditure has a long history that can be traced to the late 1800s and the industrial revolution. Machines that are run fast wear out more quickly, so the notion was born that humans and animals might experience similar fates: the faster they live (expressed as greater energy expenditure), the sooner they die. Evidence supporting the “rate-of-living” theory was gleaned from the scaling of resting metabolism and life span as functions of body mass. The product of these factors yields a mass-invariant term, equivalent to the “amount of living.” There are at least four problems with this evidence, which are summarized and reviewed in this communication: 1) life span is a poor measure of aging, 2) resting metabolism is a poor measure of energy expenditure, 3) the effects are confounded by body mass and 4) the comparisons made are not phylogenetically independent. We demonstrate that there is a poor association between resting metabolic rate (RMR) and daily energy expenditure (DEE) measured using the doubly labeled water (DLW) method at the level of species. Nevertheless, the scaling relation between DEE and body mass still has the same scaling exponent as the RMR and body mass relationship. Thus, if we use DEE rather than RMR in the analysis, the rate-of-living ideas are still supported. Data for 13 species of small mammal were obtained, where energy demands by DLW and longevity were reliably known. In these species, there was a strong negative relationship between residual longevity and residual DEE, both with the effects of body mass removed (r 0.763, F 32.1, P 0.001). Hence, the association of energy demands and life span is not attributed to the confounding effects of body size. We subjected these latter data to an analysis that extracts phylogenetically independent contrasts, and the relationship remained significant (r 0.815, F 39.74, P 0.001). Small mammals that live fast really do die young. However, there are very large differences between species in the amounts of living that each enjoy and these disparities are even greater when other taxa are included in the comparisons. Such differences are incompatible with the “rate-of-living” theory. However, the link between energetics and aging across species is reconcilable within the framework of the “free-radical damage hypothesis” and the “disposable soma hypothesis.” Within species one might anticipate the rate-of-living model would be more appropriate. We reviewed data generated from three different sources to evaluate whether this were so, studies in which metabolic rate is experimentally increased and impacts on life span followed, studies of caloric restriction and studies where links between natural variation in metabolism and life span are sought. This review reveals that there might be contrasting effects of resting and nonresting energy expenditure on aging, with increases in the former being protective and increases in the latter being harmful. J. Nutr. 132: 1583S–1597S, 2002.

Over the last 50 y there have been numerous reports of vitamin E deficiency in cats with the main cause of these deficiencies being attributed to ingestion of diets high in polyunsaturated fatty acids (PUFAs). Steatitis (“yellow fat disease”) is attributed to vitamin E deficiency and has been experimentally induced in kittens after feeding a commercial cat food with a high fish content (1) and after feeding vitamin E–deficient purified diet containing tuna oil and stripped safflower seed oil (2,3). Fish oil is a rich source of PUFAs, which are highly susceptible to oxidation, and an increased intake of fish oil has been associated with increased vitamin E requirements. Currently the minimum dietary vitamin E requirement of cats has been set at 30 IU/kg dry matter with dietary levels expected to increase 3–4 times with a diet high in PUFAs (4). To prevent vitamin E deficiency in cats fed commercially sold diets, the Association of American Feed Control Officials (AAFCO) recommends that fish oil–containing diets for cats should be supplemented with 10 IU of vitamin E for every g of fish oil per kg diet (5). However, there is no direct evidence to substantiate the latter value or the increases recommended (4,5) in adult cats. The main objective of this study was to determine the vitamin E requirements of adult cats fed high dietary levels of PUFAs from fish oil. MATERIALS AND METHODS

We studied the effects of feeding an antioxidant blend of vitamins, minerals and carotenoids to a mixed adult dog population (n = 40, mean 4.4 +/- 1.85 y) for a 16-wk period. Compared to the control group of dogs (n = 20), the antioxidant (AOX)-supplemented group of dogs (n = 20) demonstrated significant increases in plasma levels of vitamin E and taurine by 4 wk of supplementation (P < 0.01) and total antioxidant activity (as measured by ferric-reducing antioxidant power assay) by 8 wk of supplementation (P < 0.05). Following 8 wk of supplementation, the AOX-supplemented dogs also showed significant reductions in both endogenous and exogenous DNA damage (P < 0.005) compared to that of the control dogs, as measured by the comet assay. Over an 8-wk rabies vaccination course that started at 8 wk supplementation, the AOX-supplemented dogs also demonstrated significantly higher vaccine-specific virus-neutralizing antibody levels at 2, 4 and 6 wk postvaccination (P < 0.05) and a tendency toward establishing a vaccine-specific antibody response quicker than did the control group of dogs. These findings in dogs suggest that antioxidant supplementation can achieve sustained increases in circulating levels of antioxidants that exert a protective effect by a decrease in DNA damage, leading to improved immunological performance. These findings also have implications in a wider context where free-radical damage has been associated with a variety of degenerative disorders and the aging process in general.

The idea that aging should be linked to energy expenditure has a long history that can be traced to the late 1800s and the industrial revolution. Machines that are run fast wear out more quickly, so the notion was born that humans and animals might experience similar fates: the faster they live (expressed as greater energy expenditure), the sooner they die. Evidence supporting the "rate-of-living" theory was gleaned from the scaling of resting metabolism and life span as functions of body mass. The product of these factors yields a mass-invariant term, equivalent to the "amount of living." There are at least four problems with this evidence, which are summarized and reviewed in this communication: 1) life span is a poor measure of aging, 2) resting metabolism is a poor measure of energy expenditure, 3) the effects are confounded by body mass and 4) the comparisons made are not phylogenetically independent. We demonstrate that there is a poor association between resting metabolic rate (RMR) and daily energy expenditure (DEE) measured using the doubly labeled water (DLW) method at the level of species. Nevertheless, the scaling relation between DEE and body mass still has the same scaling exponent as the RMR and body mass relationship. Thus, if we use DEE rather than RMR in the analysis, the rate-of-living ideas are still supported. Data for 13 species of small mammal were obtained, where energy demands by DLW and longevity were reliably known. In these species, there was a strong negative relationship between residual longevity and residual DEE, both with the effects of body mass removed (r(2) = 0.763, F = 32.1, P < 0.001). Hence, the association of energy demands and life span is not attributed to the confounding effects of body size. We subjected these latter data to an analysis that extracts phylogenetically independent contrasts, and the relationship remained significant (r(2) = 0.815, F = 39.74, P < 0.001). Small mammals that live fast really do die young. However, there are very large differences between species in the amounts of living that each enjoy and these disparities are even greater when other taxa are included in the comparisons. Such differences are incompatible with the "rate-of-living" theory. However, the link between energetics and aging across species is reconcilable within the framework of the "free-radical damage hypothesis" and the "disposable soma hypothesis." Within species one might anticipate the rate-of-living model would be more appropriate. We reviewed data generated from three different sources to evaluate whether this were so, studies in which metabolic rate is experimentally increased and impacts on life span followed, studies of caloric restriction and studies where links between natural variation in metabolism and life span are sought. This review reveals that there might be contrasting effects of resting and nonresting energy expenditure on aging, with increases in the former being protective and increases in the latter being harmful.

Abstract Little is known about the energy requirements of pet birds. Because animals generally eat to meet energy requirements, the nutrient content of a diet must be balanced with the energy content. To formulate balanced diets for a range of bird species, both the energy needs of the relevant bird species and the energy content of the diet must be calculated. The most practical way of understanding the energy requirements of pet birds is by studying their daily energy expenditure. Factors affecting energy requirements are body size, activity patterns, environmental temperature, plumage cover, age, and physiologic state. Energy costs of maintenance are directly related to the lean body mass and the relative size of the different organs, because these are the tissues that actively use oxygen. Activity patterns also have a considerable impact; the difference between sitting and standing can affect the daily energy expenditure by as much as 42%. There are a variety of published allometric equations for determining avian energy requirements, but those derived from fasted species are inappropriate. For companion birds, equations derived for the particular body-weight range of the avian species in question are the most applicable. Two equations are recommended: one for calculating the energy requirements of avian species less than 100 g, and the second for avian species whose body weights are in the range 100–1500 g.

A series of cross-sectional and longitudinal studies conducted in humans has shown that aging is associated with a gradual decline in the maintenance energy requirement. Generally, this is equivalent to a total decrement of 20% of young adult maintenance energy requirements and is a result of a decrease in both physical activity and basal metabolic rate. Relatively few such studies have been conducted in dogs, but the results have been consistent. It appears that maintenance energy requirements decline by approximately 20%, and it is assumed that the causal factors are the same as those for humans. The situation appears to be somewhat different in cats, with evidence to date indicating that maintenance energy requirements remain constant throughout adult life. Why cats should be different from other species is not clear, but it is hypothesized that relative inactivity is typical of most cats' behavior, such that there is no obvious age-related change. In terms of feeding regimens for senior cats and dogs, it is appropriate to decrease energy provision for senior dogs by approximately 20%, whereas the energy provision for senior cats should not be decreased.

When considering the question of energy balance, it is important to take into account energy provision and the ability of aging animals to digest macronutrients and thus obtain their maintenance energy requirement. A large number of studies have been conducted in humans in an effort to establish whether aging of the gastrointestinal tract has a significant effect on availability of dietary energy. The results of these studies have been conflicting, with indications that some aspects of gut functionality do decline with age, but little evidence overall to suggest that aging humans are at risk of energy deficit due to compromised digestive efficiency. A number of digestibility studies carried out with dogs confirm that there is no evidence of an age-related decline in digestive efficiency. This knowledge makes the determination of energy provision to senior dogs relatively straightforward to calculate. Many senior cats appear to exhibit quite a marked reduction in their ability to digest macronutrients, particularly fat, efficiently. Because this reduces the overall capacity to obtain energy from the diet, it is recommended that old cats should not routinely be offered reduced energy diets. For senior cats, the feeding regimen in later life should be to offer highly digestible diets that provide as much energy as adult maintenance rations. J. Nutr. 128: 2632S–2635S, 1998.

Increasing evidence suggests involvement of free-radical species in the development of oxidative DNA damage, the consequences of which have been implicated in a number of degenerative disorders associated with the aging process. Here we report the application of a single-cell gel electrophoresis (comet) assay for assessing levels of DNA damage in canine and feline leukocytes. Leukocytes were collected from 24 healthy adult cats and dogs and subjected to DNA damage ex vivo by exposure to a range of hydrogen peroxide (H2O2) concentrations (0 -250 mol/L). The optimal concentration of H2O2 to induce a significant increase in DNA damage was 100 mol/L for both canine and feline leukocyte samples. Levels of DNA damage were assessed and quantified by visual and computer image analysis. The results obtained showed high correlations between visual scoring and computer image analysis for feline samples (percentage DNA in tail, R2 0.99; tail moment, R2 0.95; tail length, R2 0.90) and canine samples (percentage DNA in tail, R2 0.97; tail moment, R2 0.95; tail length, R2 0.91). In conclusion, this method provides a way of assessing levels of DNA damage utilizing visual and/or computer image analysis in the feline and canine systems. With the capacity of the comet assay to be able to measure end products of free-radical reactions, it is a useful tool for determining the optimal effects of dietary antioxidants on a reliable biomarker of oxidative stress such as cellular DNA status in cats and dogs. J. Nutr. 132: 1598S-1603S, 2002.

The idea that aging should be linked to energy expenditure has a long history that can be traced to the late 1800s and the industrial revolution. Machines that are run fast wear out more quickly, so the notion was born that humans and animals might experience similar fates: the faster they live (expressed as greater energy expenditure), the sooner they die. Evidence supporting the "rate-of-living" theory was gleaned from the scaling of resting metabolism and life span as functions of body mass. The product of these factors yields a mass-invariant term, equivalent to the "amount of living." There are at least four problems with this evidence, which are summarized and reviewed in this communi- cation: 1) life span is a poor measure of aging, 2) resting metabolism is a poor measure of energy expenditure, 3) the effects are confounded by body mass and 4) the comparisons made are not phylogenetically independent. We demonstrate that there is a poor association between resting metabolic rate (RMR) and daily energy expenditure (DEE) measured using the doubly labeled water (DLW) method at the level of species. Nevertheless, the scaling relation between DEE and body mass still has the same scaling exponent as the RMR and body mass relationship. Thus, if we use DEE rather than RMR in the analysis, the rate-of-living ideas are still supported. Data for 13 species of small mammal were obtained, where energy demands by DLW and longevity were reliably known. In these species, there was a strong negative relationship between residual longevity and residual DEE, both with the effects of body mass removed (r2 0.763, F 32.1, P 0.001). Hence, the association of energy demands and life span is not attributed to the confounding effects of body size. We subjected these latter data to an analysis that extracts phylogenetically indepen- dent contrasts, and the relationship remained significant (r2 0.815, F 39.74, P 0.001). Small mammals that live fast really do die young. However, there are very large differences between species in the amounts of living that each enjoy and these disparities are even greater when other taxa are included in the comparisons. Such differences are incompatible with the "rate-of-living" theory. However, the link between energetics and aging across species is reconcilable within the framework of the "free-radical damage hypothesis" and the "disposable soma hypothesis." Within species one might anticipate the rate-of-living model would be more appropriate. We reviewed data generated from three different sources to evaluate whether this were so, studies in which metabolic rate is experimentally increased and impacts on life span followed, studies of caloric restriction and studies where links between natural variation in metabolism and life span are sought. This review reveals that there might be contrasting effects of resting and nonresting energy expenditure on aging, with increases in the former being protective and increases in the latter being harmful. J. Nutr. 132: 1583S-1597S, 2002.