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Using Gross Energy Improves Metabolizable Energy Predictive Equations for Pet Foods Whereas Undigested Protein and Fiber Content Predict Stool Quality
Open accessAbstract
Because animal studies are labor intensive, predictive equations are used extensively for calculating metabolizable energy (ME) concentrations of dog and cat pet foods. The objective of this retrospective review of digestibility studies, which were conducted over a 7-year period and based upon Association of American Feed Control Officials (AAFCO) feeding protocols, was to compare the accuracy and precision of equations developed from these animal feeding studies to commonly used predictive equations. Feeding studies in dogs and cats (331 and 227 studies, respectively) showed that equations using modified Atwater factors accurately predict ME concentrations in dog and cat pet foods (r²= 0.97 and 0.98, respectively). The National Research Council (NRC) equations also accurately predicted ME concentrations in pet foods (r² = 0.97 for dog and cat foods). For dogs, these equations resulted in an average estimate of ME within 0.16% and 2.24% of the actual ME measured (equations using modified Atwater factors and NRC equations, respectively); for cats these equations resulted in an average estimate of ME within 1.57% and 1.80% of the actual ME measured. However, better predictions of dietary ME in dog and cat pet foods were achieved using equations based on analysis of gross energy (GE) and new factors for moisture, protein, fat and fiber. When this was done there was less than 0.01% difference between the measured ME and the average predicted ME (r² = 0.99 and 1.00 in dogs and cats, respectively) whereas the absolute value of the difference between measured and predicted was reduced by approximately 50% in dogs and 60% in cats. Stool quality, which was measured by stool score, was influenced positively when dietary protein digestibility was high and fiber digestibility was low. In conclusion, using GE improves predictive equations for ME content of dog and cat pet foods. Nondigestible protein and fiber content of diets predicts stool quality.
Overview
CoChat Summary
TL;DR
In a retrospective review of 558 AAFCO-based digestibility studies, predictive equations for metabolizable energy (ME) in dog and cat foods were accurate when using modified Atwater or NRC factors, but equations incorporating measured gross energy (GE) plus moisture, protein, fat, and fiber improved accuracy and precision. Stool quality was better when less undigested protein and more fiber reached the large intestine.
Key findings
- Modified Atwater factors predicted ME accurately in dog foods (r² = 0.97) and cat foods (r² = 0.98); NRC equations also gave r² = 0.97 for both dogs and cats. Average predicted ME differed from measured ME by 0.16% and 2.24% in dogs using modified Atwater and NRC equations, respectively, and by 1.57% and 1.80% in cats.
- New equations using measured GE and new coefficients for moisture, protein, fat, and fiber yielded r² = 0.99 in dogs and 1.00 in cats, with less than 0.01% difference between average measured and predicted ME; the absolute difference between measured and predicted ME was reduced by approximately 50% in dogs and 60% in cats.
- In dogs, stool score was higher when less protein entered the large intestine (4.3 ± 0.04 vs. 4.1 ± 0.04, P≤0.05) and when more fiber entered the large intestine (4.2 ± 0.04 vs. 4.1 ± 0.04, P≤0.05). In cats, stool score was higher when less protein entered the large intestine (4.1 ± 0.05 vs. 3.7 ± 0.05, P≤0.05) and when more fiber entered the large intestine (4.0 ± 0.07 vs. 3.8 ± 0.06, P≤0.05).
- There was no correlation between dietary protein concentration and stool quality.
Methods
- Retrospective review of 558 digestibility studies conducted over 7 years: 331 studies in healthy adult Beagle dogs (124 dogs; mean age 6.7 years, range 2–12) and 227 studies in healthy adult domestic short hair cats (138 cats; mean age 8.0 years, range 1–15). Each test used six adult dogs or cats and followed the AAFCO quantitative collection protocol, with a pre-collection phase of at least 7 days and a 5-day (120-hour) total fecal collection phase.
- Foods included dry and canned dog and cat foods, both commercial and non-commercial, all meeting AAFCO requirements for complete and balanced adult dog or cat foods. Food energy, moisture, protein, fat, fiber, and ash were analyzed by a commercial laboratory using AOAC methods; ME was measured with AAFCO methods and compared with modified Atwater, NRC, and new study-derived predictive equations.
- Statistical analyses used SAS general linear models; multivariate regression identified equations minimizing variation between estimated and actual ME. Stool quality was scored on a 1–5 scale by trained, masked graders. Nutrient digestibility effects on stool quality were evaluated with species and form (dry or canned) as class variables and measured dietary ME as a covariate; high/low classifications for protein or fiber entering the large intestine were based on median split, and mean separation used Duncan’s least significant difference test with significance at P≤0.05.
Limitations
- The paper does not state an explicit limitations section.
- Evident from the text: the new ME equations were derived from and evaluated against the same set of 558 digestibility studies, with no independent validation reported, so the high accuracy and precision may be optimistic. The studies were conducted at a single commercial sponsor (Hill’s Pet Nutrition) using animals owned by the funders, which may limit generalizability.
- Evident from the text: stool quality relied on subjective 1–5 grading, though graders were trained, previously evaluated, and masked to foods. The study population was healthy adult dogs and cats fed AAFCO-complete foods, so findings may not extend to other ages, health states, or non-AAFCO diets.
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Figures
![Figure 1 Relationship between measured metabolizable energy (ME) concentrations (x-axis) and ME concentrations predicted using equations with modified Atwater factors [4] (y-axis) for dog and cat pet foods. The modified Atwater factors are 3.5 kcal/g for protein and carbohydrate, and 8.5 kcal/g for fat. Ideally, all points should be on the line x = y. A) Measured ME concentrations were determined from 331 total digestibility studies in dogs, of which 259 used dry dog foods and 72 used canned dog foods. B) Measured ME concentrations were determined from 227 total digestibility studies in cats, of which 173 used dry cat foods and 54 used canned cat foods.](/paper-figures/2f49445baa7e49b8a095d11142c6871a/using-gross-energy-improves-metabolizable-energy-predictive-equations-for-pet-figure-1.jpg)
![Figure 2 Relationship between measured metabolizable energy (ME) concentrations (x-axis) and ME concentrations predicted using National Research Council (NRC) equations [2] (y-axis) for dog and pet foods. Ideally, all points should be on the line x = y. A) Measured ME concentrations were determined from 331 total digestibility studies in dogs, of which 259 used dry dog foods and 72 used canned dog foods. The NRC equations for dogs first calculate gross energy (GE) using the equation GE = 5.7× g protein +9.4× g fat +4.1× (g NFE + g fiber). Energy digestibility coefficients are then calculated for dogs as (91.2–1.43× percentage crude fiber in DM). These digestibility coefficients then allow calculation of digestible energy (DE) in dogs as DE = GE × percentage energy digestibility/100 and, subsequent calculation of ME as ME = DE – (1.04× g protein). B) Measured ME concentrations were determined from 227 total digestibility studies in cats, of which 173 used dry cat foods and 54 used canned cat foods. The NRC equations for cats first calculate GE using the equation GE = 5.7× g protein +9.4× g fat +4.1× (g NFE + g fiber). Energy digestibility coefficients are then calculated for cats as (87.9–0.88× percentage crude fiber in DM). These digestibility coefficients then allow calculation of DE in cats as DE = GE × percentage energy digestibility/100 and, subsequent calculation of ME as ME = DE – (0.77× g protein).](/paper-figures/7e185da7d7fb46b9a8fa89be97e8f208/using-gross-energy-improves-metabolizable-energy-predictive-equations-for-pet-figure-2.jpg)

Verification
Claims the paper makes
0 Valid · 0 Not valid · 20 Not checked · 42 Own results and interpretations
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“The National Research Council (NRC) equations also accurately predicted ME concentrations in pet foods (r2 = 0.97 for dog and cat foods).”
§ Using Gross Energy Improves Metabolizable Energy Predictive Equations for Pet Foods Whereas Undigested Protein and Fiber Content Predict Stool Quality > Abstract
plainstates an association- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- Rests on an assumption not shown here.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Stool quality, which was measured by stool score, was influenced positively when dietary protein digestibility was high and fiber digestibility was low.”
§ Using Gross Energy Improves Metabolizable Energy Predictive Equations for Pet Foods Whereas Undigested Protein and Fiber Content Predict Stool Quality > Abstract
plainstates a cause- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“for cats these equations resulted in an average estimate of ME within 1.57% and 1.80% of the actual ME measured.”
§ Using Gross Energy Improves Metabolizable Energy Predictive Equations for Pet Foods Whereas Undigested Protein and Fiber Content Predict Stool Quality > Abstract
plainstates an association- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- Rests on an assumption not shown here.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“For dogs, these equations resulted in an average estimate of ME within 0.16% and 2.24% of the actual ME measured (equations using modified Atwater factors and NRC equations, respectively)”
§ Using Gross Energy Improves Metabolizable Energy Predictive Equations for Pet Foods Whereas Undigested Protein and Fiber Content Predict Stool Quality > Abstract
plainstates an association- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- Rests on an assumption not shown here.
A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“However, better predictions of dietary ME in dog and cat pet foods were achieved using equations based on analysis of gross energy (GE) and new factors for moisture, protein, fat and fiber.”
§ Using Gross Energy Improves Metabolizable Energy Predictive Equations for Pet Foods Whereas Undigested Protein and Fiber Content Predict Stool Quality > Abstract
plainstates a cause- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- Rests on an assumption not shown here.
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Citations
Cites
Small animal clinical nutrition
Hand, M. S., Morris, M. L., & Novotny, B. J. (2000) · Medical Entomology and Zoology · Cited by 273
Lubbs, D. C., Vester, B. M., Fastinger, N. D., & Swanson, K. S. (2009) · Journal of Animal Physiology and Animal Nutrition · doi:10.1111/j.1439-0396.2007.00788.x · Cited by 103
Hang, I., Rinttila, T., Zentek, J., Kettunen, A., Alaja, S., Apajalahti, J., Harmoinen, J., de Vos, W. M., & Spillmann, T. (2012) · BMC Veterinary Research · doi:10.1186/1746-6148-8-90 · Cited by 75
Lewis, L. D., Magerkurth, J. H., Roudebush, P., Morris, M. L., Mitchell, E. E., & Teeter, S. M. (1994) · The Journal of Nutrition · doi:10.1093/jn/124.suppl_12.2716s · Cited by 39
Zentek, J., Fricke, S., Hewicker-Trautwein, M., Ehinger, B., Amtsberg, G., & Baums, C. (2004) · The Journal of Nutrition · doi:10.1093/jn/134.8.2158s · Cited by 38
Determining metabolizable energy content in commercial pet foods
Laflamme, D. P. (2001) · Journal of Animal Physiology and Animal Nutrition · doi:10.1046/j.1439-0396.2001.00330.x · Cited by 34
Prola, L., Dobenecker, B., Mussa, P. P., & Kienzle, E. (2010) · Journal of Animal Physiology and Animal Nutrition · doi:10.1111/j.1439-0396.2008.00916.x · Cited by 33
Effect of cellulose on the digestibility of high starch versus high fat diets in dogs
Kienzle, E., Dobenecker, B., & Eber, S. (2001) · Journal of Animal Physiology and Animal Nutrition · doi:10.1046/j.1439-0396.2001.00315.x · Cited by 31
Influence of different cellulose types on feces quality of dogs
Wichert, B., Schuster, S., Hofmann, M., Dobenecker, B., & Kienzle, E. (2002) · The Journal of Nutrition · doi:10.1093/jn/132.6.1728s · Cited by 29
Further developments in the prediction of metabolizable energy (ME) in pet food
Kienzle, E. (2002) · The Journal of Nutrition · doi:10.1093/jn/132.6.1796s · Cited by 18
Research works printed but not linked: 3
- NRC (USA) (2006) Nutrient requirements of dogs and cats/Ad Hoc Committee on Dog and Cat Nutrition. Natl Acad Press, Washington, DC.
- AAFCO (2012) Association of American Feed Control Officials. Oxford, IN: Official Publication.
- PT Kendall, 1985, Methods of estimation of the metabolizable energy content of cat foods, Feline Practice, 15, 38
Cited by
Hall, J. A., Yerramilli, M., Obare, E., Yerramilli, M., Yu, S., & Jewell, D. E. (2014) · The Veterinary Journal · doi:10.1016/j.tvjl.2014.10.021 · Cited by 123
Self-citationJackson, M. I., & Jewell, D. E. (2018) · Gut Microbes · doi:10.1080/19490976.2018.1526580 · Cited by 98
Self-citationVarying Protein Levels Influence Metabolomics and the Gut Microbiome in Healthy Adult Dogs
Ephraim, E., Cochrane, C., & Jewell, D. E. (2020) · Toxins · doi:10.3390/toxins12080517 · Cited by 72
Self-citationHall, J. A., Jackson, M. I., Jewell, D. E., & Ephraim, E. (2020) · PLoS ONE · doi:10.1371/journal.pone.0235480 · Cited by 58
Self-citationHall, J. A., MacLeay, J. M., Yerramilli, M., Obare, E., Yerramilli, M., Schiefelbein, H. M., Paetau‐Robinson, I., & Jewell, D. E. (2016) · PLoS ONE · doi:10.1371/journal.pone.0153654 · Cited by 50
Self-citation
Metadata
- Authors
- Hall, J. A., Melendez, L. D., & Jewell, D. E.
- Affiliations
- Oregon State University (US)
- Hill's Pet Nutrition (United States) (US)
- Corresponding author
- Hall, Jean A.
- Venue
- PLoS ONE
- Publisher
- Public Library of Science (PLoS)
- Published
- 2013-01-14
- Volume
- 8
- Issue
- 1
- Pages
- e54405
- ISSN
- 1932-6203 (ISSN-L)
- PMC
- PMC3544805
- OpenAlex
- W2084309644
- Cited by
- 92
- References
- 17
- Citations by year
- 2026: 12 · 2025: 9 · 2024: 10 · 2023: 10 · 2022: 12 · 2021: 7 · 2020: 12 · 2019: 3 · 2018: 3 · 2017: 5 · 2016: 3 · 2015: 3 · 2014: 2
- FWCI
- 3.95
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Locations
- PLoS ONE (publishedVersion, cc-by)
- PubMed (publishedVersion)
- PubMed Central (publishedVersion, cc-by)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- Funders
- Hill's Pet Nutrition
- Topics
- Life Sciences › Agricultural and Biological Sciences › Animal Science and Zoology › Animal Nutrition and Physiology
- Health Sciences › Veterinary › Small Animals › Veterinary Medicine and Surgery
- Life Sciences › Agricultural and Biological Sciences › Aquatic Science › Aquaculture Nutrition and Growth
- Keywords
- metabolizable energy, gross energy, prediction equations, dog food, cat food, pet food, Atwater factors, diet digestibility, protein digestibility, fiber digestibility, stool quality
- MeSH terms
- Animals, Animal Feed, Cats, Dietary Fiber, Digestion, Dogs, Feces, Nutritive Value, Retrospective Studies
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