Richard F. Butterwick

Validation of dual energy X-ray absorptiometry (DXA) by comparison with chemical analysis of dogs and cats

BACKGROUND: Dual-energy X-ray absorptiometry (DXA) has been used extensively to measure body composition, but has been validated by comparison to chemical analysis on relatively few occasions. Moreover, these previous validation studies have ground up entire carcasses prior to chemical analysis, thus potentially obscuring sources of error in the DXA analysis.OBJECTIVE: The purpose of this study was to validate DXA by comparison to chemical analysis in dogs and cats, performing chemical analysis on dissected rather than ground carcasses to reveal sources of discrepancy between the two methods.DESIGN: Sixteen animals (10 cats and 6 dogs weighing between 1.8 and 22.1 kg) were scanned by DXA post-mortem using a Hologic QDR-1000 W pencil beam machine and then dissected into 22 separate components. Individual tissues were dried and then sub-sampled for analysis of fat content by Soxholet extraction, or ashing in a muffle furnace. Body composition by DXA was compared to body composition by chemical analysis and discrepancies between the two correlated with chemical composition of individual tissues. We also explored the capability of the machine to establish the fat contents of mixtures of ground beef, lard and water.RESULTS: DXA estimates were strongly correlated with estimates derived from chemical analysis: total body mass (r=1.00), lean tissue mass (r=0.999), body water content (r=0.992) and fat mass (r=0.982). Across individuals the absolute and percentage discrepancies were also small: total body mass (13.2 g, 1.02%), lean tissue mass (119.4 g, 2.64%), water content (101 g, 1.57%) and fat content (28.5 g, 2.04%), where the percentage error is expressed relative to the average mass of that component across all individuals. Although on average DXA compared very well to chemical analysis, individual errors were much greater. Individual errors in the lean tissue and fat tissue components were strongly correlated with the fat content of skeletal muscle and the lean content of mesenteric fat. The error in the DXA estimate of total fat content was related to skeletal muscle hydration. Experimental studies using mixtures of lean ground beef, water and lard indicated that tissue hydration may have important effects on the perception of tissue fat content by DXA. Bone mineral content by DXA was approximately 30% lower than whole body ash content but only 7.7% lower than ash content of the bones.CONCLUSIONS: On average pencil beam DXA analysis using the Hologic QDR-1000 W machine provides an accurate estimate of body composition in subjects weighing between 1.8 and 22.1 kg. Individual discrepancies, however, can be large and appear to be related to lean tissue hydration.

Geometric analysis of macronutrient selection in breeds of the domestic dog, Canis lupus familiaris.

Although many herbivores and omnivores have been shown to balance their intake of macronutrients when faced with nutritionally variable foods, study of this ability has been relatively neglected in carnivores, largely on the assumption that prey are less variable in nutrient composition than the foods of herbivores and omnivores and such mechanisms therefore unnecessary. We performed diet selection studies in 5 breeds of adult dog (Canis lupus familiaris) to determine whether these domesticated carnivores regulate macronutrient intake. Using nutritional geometry, we show that the macronutrient content of the diet was regulated to a protein:fat:carbohydrate ratio of approximately 30%:63%:7% by energy, a value that was remarkably similar across breeds. These values, which the analysis suggests are dietary target values, are based on intakes of dogs with prior experience of the respective experimental food combinations. On initial exposure to the diets (i.e., when naive), the same dogs self-selected a diet that was marginally but significantly lower in fat, suggesting that learning played a role in macronutrient regulation. In contrast with the tight regulation of macronutrient ratios, the total amount of food and energy eaten was far higher than expected based on calculated maintenance energy requirements. We interpret these results in relation to the evolutionary history of domestic dogs and compare them to equivalent studies on domestic cats.

Advances in Dietary Management of Obesity in Dogs and Cats

Recent evidence in humans has reemphasized the importance of specific lifestyle behaviors such as activity level on energy requirements. A recent survey of adult pet dogs has shown a clear association between the level of activity and energy requirement, and suggests that current feeding recommendations may overestimate the energy requirements of adult dogs. Although a reduction in feeding guides may help to reduce the risk of overfeeding and subsequent development of obesity in adult dogs, there is considerable individual variation in energy requirements, which emphasizes the importance of tailoring feeding practices to the individual. Diet clearly has a critical role to play in both the prevention and treatment of obesity. We have evaluated the effect of different dietary regimens on the treatment of obesity in companion animals. In cats, increased energy restriction results in more rapid weight loss. However, this is associated with less favorable changes in body composition. In dogs, we have evaluated the potential benefit of insoluble and soluble dietary fiber on satiety in dogs that have been restricted to an energy intake appropriate for weight reduction. Results of a series of studies have failed to show any benefit of either fiber type on satiety in energy-restricted dogs.

Energetics of lactation in domestic dog (Canis familiaris) breeds of two sizes

The energetics of lactation was measured in two breeds of domestic dog during peak lactation. Labrador Retrievers (30 kg) had larger litter sizes than Miniature Schnauzers (6 kg). During the 7-day experimental period, Labrador pups increased more in mass than Schnauzer pups, both absolutely and relatively. Consequently, the energy demands of the litter, relative to maternal metabolism, were higher in Labradors than Schnauzers. Milk composition and gross efficiency of milk production were not significantly different between breeds and the costs of lactation were fuelled by increases in food intake. Metabolisable energy intake was higher than predicted in Labradors, but lower than predicted in Schnauzers. These patterns differ from interspecific expectations, which would predict larger animals to reproduce more slowly, have smaller litter sizes, and invest less energy in reproduction.

The effect of dietary starch level on postprandial glucose and insulin concentrations in cats and dogs

A charge made against feeding dry foods to cats is that the high carbohydrate (i.e. starch) content results in high blood glucose levels which over time may have detrimental health effects. The present study determined the post-meal concentrations of plasma glucose and insulin in adult cats (seven males and four females) and dogs (Labrador retrievers; four males and five females) fed dry diets with low-starch (LS), moderate-starch (MS) or high-starch (HS) levels. In a cross-over design with at least 7 d between the test meals, plasma glucose and insulin concentrations were measured following a single meal of a LS, MS and HS diet (209 kJ/kg bodyweight). Only the HS diet resulted in significant post-meal increases in plasma glucose concentration in cats and dogs although the time-course profiles were different between the species. In cats, plasma glucose concentration was significantly increased above the pre-meal concentration from 11 h until 19 h after the meal, while in dogs, a significant increase above baseline was seen only at the 7 h time point. Plasma insulin was significantly elevated in dogs 4-8 h following the MS diet and 2-8 h after the HS diet. In cats, plasma insulin was significantly greater than baseline from 3-7 and 11-17 h after the HS diet. The time lag (approximately 11 h) between eating the HS diet and the subsequent prolonged elevation of plasma glucose concentration seen in cats may reflect metabolic adaptations that result in a slower digestive and absorptive capacity for complex carbohydrate.

Postprandial glucose and insulin profiles following a glucose-loaded meal in cats and dogs

Data from intravenous (i.v.) glucose tolerance tests suggest that glucose clearance from the blood is slower in cats than in dogs. Since different physiological pathways are activated following oral administration compared with i.v. administration, we investigated the profiles of plasma glucose and insulin in cats and dogs following ingestion of a test meal with or without glucose. Adult male and female cats and dogs were fed either a high-protein (HP) test meal (15 g/kg body weight; ten cats and eleven dogs) or a HP + glucose test meal (13 g/kg body-weight HP diet + 2 g/kg body-weight D-glucose; seven cats and thirteen dogs) following a 24 h fast. Marked differences in plasma glucose and insulin profiles were observed in cats and dogs following ingestion of the glucose-loaded meal. In cats, mean plasma glucose concentration reached a peak at 120 min (10.2, 95 % CI 9.7, 10.8 mmol/l) and returned to baseline by 240 min, but no statistically significant change in plasma insulin concentration was observed. In dogs, mean plasma glucose concentration reached a peak at 60 min (6.3, 95 % CI 5.9, 6.7 mmol/l) and returned to baseline by 90 min, while plasma insulin concentration was significantly higher than pre-meal values from 30 to 120 min following the glucose-loaded meal. These results indicate that cats are not as efficient as dogs at rapidly decreasing high blood glucose levels and are consistent with a known metabolic adaptation of cats, namely a lack of glucokinase, which is important for both insulin secretion and glucose uptake from the blood.

Effect of Level and Source of Dietary Fiber on Food Intake in the Dog

The effects of dietary fiber on challenge meal intake and on the perception of hunger in dogs were evaluated. A program of testing variants of a standard low energy diet, to which one of five fiber containing raw materials was added, was undertaken. Diets were fed to a group of six dogs for 12-d periods in a latin square design and in amounts that corresponded to the food allowance for weight reduction. Behavioral characteristics of dogs were recorded on videotape for 30-min periods after introduction of test diets. On two occasions during each 12-d feeding period dogs were presented with a challenge meal. At the end of each 12-d feeding period all dogs entered a 6-d washout period. There was no significant effect of diet on the intake of the challenge meal or on intake of food during the subsequent washout period. In addition, diet had no apparent effect on the perception of hunger, as represented by behavioral characteristics during the 30-min period after presentation of test diets. It was concluded that inclusion of moderate levels of raw materials, composed primarily of insoluble fiber, in a commercial low energy diet had no apparent beneficial effects on satiety, when fed to dogs on an energy intake corresponding to allowances for weight reduction.

Effects of neutering on food intake, body weight and body composition in growing female kittens

To understand the effects of neutering on food intake, body weight (BW) and body composition in kittens, data from an unrelated study were subjected to post hoc analysis. A total of twelve pairs of 11-week-old female littermates were randomly assigned to either a neutered group (neutered at 19 weeks old) or an entire group (kept entire) and offered free access to a dry diet until the age of 1 year. Neutered kittens exhibited increased food intake and increased BW after neutering (both P < 0.00 001). Food intake (per kg BW) peaked 10 weeks after neutering; the mean intake of neutered kittens was 17 (95 % CI 8, 27) % more than entire littermates (P = 0.00 014). The intake was then reduced until there was no significant difference between the groups 18 weeks post-neutering. By 52 weeks of age, the neutered kittens were 24 (95 % CI 11, 39) % heavier than entire littermates (P < 0.0001) with a body condition score (BCS) 16.6 (95 % CI 0.9, 34.8) % higher (P = 0.0028). Neutered kittens continued to grow significantly fatter after neutering (all P < 0.0014), while entire kittens showed no significant change after 18 weeks of age. As neutered kittens consumed similar amounts of energy to their entire littermates from 18 weeks post-neutering, while their BW, BCS and percentage fat continued to increase, we suggest that neutered kittens have a reduced metabolisable energy requirement, and should therefore be fed to maintain an ideal BCS rather than ad libitum. Moreover, to maintain an ideal BCS, entire kittens consumed 93 (95 % CI 87, 100) % of their theoretical intake at 26 weeks of age, and 79 (95 % CI 72, 87) % at 52 weeks of age, suggesting that the current energy recommendation is inappropriate for these kittens.

Energy Requirements of Adult Dogs: A Meta-Analysis

A meta-analysis was conducted to determine the maintenance energy requirements of adult dogs. Suitable publications were first identified, and then used to generate relationships amongst energy requirements, husbandry, activity level, methodology, sex, neuter status, dog size, and age in healthy adult dogs. Allometric equations for maintenance energy requirements were determined using log-log linear regression. So that the resulting equations could readily be compared with equations reported by the National Research Council, maintenance energy requirements in the current study were determined in kcal/kg0.75 body weight (BW). Ultimately, the data of 70 treatment groups from 29 publications were used, and mean (± standard deviation) maintenance energy requirements were 142.8±55.3 kcal.kgBW−0.75.day−1. The corresponding allometric equation was 81.5 kcal.kgBW−0.93.day−1 (adjusted R2 = 0.64; 70 treatment groups). Type of husbandry had a significant effect on maintenance energy requirements (P<0.001): requirements were greatest in racing dogs, followed by working dogs and hunting dogs, whilst the energy requirements of pet dogs and kennel dogs were least. Maintenance energy requirements were less in neutered compared with sexually intact dogs (P<0.001), but there was no effect of sex. Further, reported activity level tended to effect the maintenance energy requirement of the dog (P = 0.09). This review suggests that estimating maintenance energy requirements based on BW alone may not be accurate, but that predictions that factor in husbandry, neuter status and, possibly, activity level might be superior. Additionally, more information on the nutrient requirements of older dogs, and those at the extremes of body size (i.e. giant and toy breeds) is needed.

Energetics and litter size variation in domestic dog Canis familiaris breeds of two sizes

We measured resting metabolic rate (RMR), daily energy expenditure (DEE) and metabolisable energy intake (MEI) in two breeds of dog during peak lactation to test whether litter size differences were a likely consequence of allometric variation in energetics. RMR of Labrador retrievers (30 kg, n=12) and miniature Schnauzers (6 kg, n=4) averaged 3437 and 1062 kJ/day, respectively. DEE of Labradors (n=6) and Schnauzers (n=4) averaged 9808 and 2619 kJ/day, respectively. MEI of Labradors (n=12) was 22448 kJ/day and of Schnauzers (n=7) was 5382 kJ/day. DEE of Labrador pups (2.13 kg, n=19) was 974 kJ/day and Schnauzers (0.89 kg, n=7) were 490 kJ/day. Although Labradors had higher MEIs than Schnauzers during peak lactation, there was no difference in mass-specific energy expenditure between the two breeds. Hence, it is unlikely that litter size variation is a likely consequence of differences in maternal energy expenditure. Individual offspring were relatively more costly for mothers of the smaller breed to produce. Therefore, litter size variations were consistent with the expectation that smaller offspring should be more costly for mothers, but not that smaller mothers should per se invest more resources in reproduction.