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Overview The National Research Council has published several chapter utilizes older well-established literature as well as editions of Nutrient Requirements of Dogs and of Nutrient the new literature published since the last editions of Requirements of Cats, with the most recent editions pub- Nutrient Requirements of Dogs (1985) and Nutrient lished in 1985 and 1986, respectively. This new volume rep- Requirements of Cats (1986). As the committee delved into resents a revision of both of the previous reports. its task, it became apparent that the committee needed to Nutrient Requirements of Dogs and Cats has used the lat- examine earlier research as well to formalize the boundaries est scientific information to provide the requirements for of knowledge for each topic explored in this publication. In individual nutrients and the scientific basis for the require- some instances, such as digestive physiology and physical ments of healthy dogs and cats at several stages of growth activity, the chapters are essentially new additions requiring and physiological states. The specific nutrients required for a thorough review of literature through many decades. normal growth and development and their estimated daily The committee was also cognizant of the various audi- needs have been elucidated in previous editions of the ences for this report. The report will be used by profession- Nutrient Requirements series. This edition contains the lat- als in industry and academia for formulating diets and iden- est data on requirements that are based on the utilization of tifying new topics for research. Government officials may nutrients in ingredients commonly produced and commer- use the report as guidance for regulations for pet food label- cially available in dog and cat foods rather than only on ing. Students and teachers at universities will use the report purified diets. Because of overlapping use of feed ingredi- as a textbook for dog and cat nutrition. Finally, pet owners ents, as a convenience to readers and for efficient presenta- will use the report in evaluating the feeding decisions for tion of the scientific basis for nutrient requirements, the their pets. With these varied audiences, the committee chose Committee on Animal Nutrition recommended combining to err on the side of caution and include adequate detail from the dog and cat data that previously had been published sep- the literature cited to provide a clear roadmap for how the arately (National Research Council, 1985, 1986). recommendations were derived. Nutrient requirement data presented in this report are derived from peer-reviewed literature. An extensive amount MINIMAL REQUIREMENT, ADEQUATE INTAKE, of new research conducted since the previous National RECOMMENDED ALLOWANCE, AND SAFE Research Council publications on dogs and cats was avail- UPPER LIMIT able for this NRC report, yet several gaps still exist in our knowledge of requirements for specific nutrients. These Data on the daily provision of nutrients to dogs and cats gaps in our knowledge are noted in the text and by the are presented in tabular form at the end of this publication. absence of data in the requirement tables. Tables of ingredi- First, the Minimal Requirement is presented and is defined ent composition are provided so that users can select the as the minimal concentration or amount of a bioavailable most common ingredients for the development of diets that nutrient that will support a defined physiological state. Data provide adequate intake of the required nutrients for optimal on Adequate Intake follow. Adequate Intake is defined as the growth, maintenance, gestation, and lactation of dogs and concentration in the diet or amount required by the animal cats. of a nutrient that is presumed to sustain a given life stage The report is organized into 15 chapters, each addressing when no Minimal Requirement has been demonstrated. unique aspects of nutrition of dogs and cats, or providing Next, the Recommended Allowance for each nutrient is pre- summary tables of requirements and feed ingredients. Each sented. The Recommended Allowance is defined as the con- 1

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2 NUTRIENT REQUIREMENTS OF DOGS AND CATS centration or amount of a nutrient in a diet formulated to the majority of data were obtained from dogs between 5 and support a given physiological state. The Recommended 25 kg BW. The equations for energy requirements in cats are Allowance is based on the Minimal Requirement and, where based on data of domestic cats between 2.5 and 7 kg of applicable, includes a bioavailability factor. If no Minimal mature weight. Energy requirements for cats at maintenance Requirement is available, the Recommended Allowance is are expressed as kcal of ME per kilogram BW0.67 per day for based on the Adequate Intake. Last, data for Safe Upper cats with body condition scores equal to or less than 5.0 (on Limit, or the maximal concentration or amount of a nutrient a 9 point scale) and per kilogram BW0.4 for cats with body that has not been associated with adverse effects, are scores greater than 5.0. Energy requirements for pregnant presented. queens are expressed as a function of BW0.67, whereas for As noted in the footnotes to the requirement tables, the lactating queens, energy requirements are a function of both following assumptions were made to standardize the tabu- BW0.67 and BW as well as number of suckling kittens. Daily lated data. The energy density of the diet for both dogs and energy requirements for growing cats are based on mainte- cats was assumed to be 4,000 kilocalories (kcal) of metabo- nance requirements multiplied by a factor that is a function lizable energy (ME) per kilogram (kg). Requirements for of the ratio between actual BW and expected mature BW. growth of puppies are based on a 5.5-kg puppy that con- sumes 1,000 kcal of ME per day. Requirements for adult REPORT HIGHLIGHTS dogs at maintenance are based on a 15-kg adult dog that consumes 1,000 kcal of ME per day. Requirements for ges- Digestive physiology is examined in Chapter 1. The tating and lactating dogs are based on a 22-kg bitch with chapter, which was not previously published in the dog and eight puppies in peak lactation consuming 5,000 kcal of ME cat series, includes three components: (1) digestive com- per day. Requirements for growth of kittens are based on an partments and functions; (2) hormonal aspects of digestion; 800-g kitten consuming 180 kcal of ME per day, and those and (3) measurement of, and factors affecting, digestibility. for an adult cat at maintenance are based on a 4-kg adult cat This description of each species sets the stage for under- consuming 250 kcal of ME per day. Requirements for ges- standing digestive function and nutritional needs of dogs tating and lactating cats are based on 4-kg queen with four and cats. kittens in peak lactation consuming 540 kcal of ME per day. Chapter 2 addresses feeding behavior of dogs and cats. For each of the previously mentioned categories of daily This new chapter examines intake and factors that affect nutrient requirement data, units of expression are, in order, food and water intake, including aspects of feeding behavior (1) amount of nutrient per kilogram of dietary dry matter and ingestion that are different in dogs and cats. The chap- (DM), which is assumed to contain 4,000 kcal, (2) amount ter includes discussions of how diurnal rhythms, social of nutrient per 1,000 kcal of ME, and (3) amount of nutrient behavior, dietary deficiencies, and learned taste aversions per kilogram of body weight to the 0.75 power (BW0.75) for are involved in feeding behavior such as quantity of food dogs, and per kilogram of body weight to the 0.67 power eaten, palatability, and dietary choice. (BW0.67) for cats. Expressing a requirement relative to DM A discussion of specific nutrients begins in Chapter 3 or ME is most convenient when formulating a diet, but with energy. The chapter examines ways to predict energy expressing a requirement relative to metabolic body weight content of diets. The basis for energy requirement recom- may be more convenient when formulating a diet for an mendations is explored thoroughly through discussions on individual dog or cat. Unfortunately, requirements expressed the basal metabolic rate, diet-induced thermogenesis, and relative to DM change with the energy density of the diet, resting fed metabolic rate for both dogs and cats, leading to and, in some instances, requirements expressed relative to energy requirement recommendations for maintenance, ME may change with body weight (BW). These changes are growth, pregnancy, and lactation. The chapter concludes illustrated in the introduction to the nutrient requirement with a discussion of energy deficiency and excess including tables in Chapter 15. body condition score evaluation. Energy requirements are expressed as kcal of ME per The discussion of energy is followed by a detailed kilogram BW0.75 per day for dogs at maintenance and for description of carbohydrates and fiber in dog and cat nutri- pregnant bitches. For lactating bitches, energy requirements tion (Chapter 4). This discussion is organized around four are a function of both BW0.75 and BW as well as number of types of carbohydrates: (1) absorbable, (2) digestible, suckling puppies. Daily energy requirements for growing (3) fermentable, and (4) nonfermentable carbohydrates. dogs are based on maintenance requirements multiplied by Digestion, absorption, utilization, and nutritive value are a factor that is a function of the ratio between actual BW and explored for each type of carbohydrate. The use of carbohy- expected mature BW. The equations for energy require- drates in dog and cat diet formulations ends the chapter. ments in dogs are based on data of dogs between 4 and 60 Chapter 5 addresses fat and fatty acid (lipid) nutrition kg of mature weight, except for female reproduction where with emphasis on the metabolism of the various fatty acid

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OVERVIEW 3 types. This chapter has been updated and expanded to Special considerations for the nutrition and feeding of dogs include nomenclature, types and sources of dietary fats, ana- and cats as laboratory animals are discussed in Chapter 10. lytical procedures, digestion/absorption of fats, and their Entirely new to this combined revision are the effects of digestibility. It also examines the biochemical basis of fatty physical activity and the environment on nutrient require- acid essentiality, describes desaturation and chain elonga- ments of dogs and cats in Chapter 11. Starting with dogs as tion pathways, and discusses both structural and functional athletes (sled and racing dogs), effects of sprinting and aspects of dietary lipids. In addition, summaries of studies endurances activities on nutrient requirements are explored. specifically conducted in dogs and cats with mention of The effects of temperature (both high and low) and high important comparative studies common to other mammalian altitude (lower oxygen supply) are examined for energy species also are included. Based on this information, suffi- requirements. Nutrient requirements as a function of cient data are now available to better address the essential amount of exercise and ambient temperatures are dis- nature of both n-6 and n-3 fatty acid types. The chapter cul- cussed for water, protein, fats, carbohydrates, minerals, and minates with an up-to-date discussion of the requirements, vitamins. recommendations, and allowances for dogs and cats at their The committee presents concepts of diet formulation in various recognized life stages. Chapter 12. Even though substantial differences in the nutri- Chapter 6 addresses protein and amino acid nutrition, ent requirements of dogs and cats exist, there are many sim- with emphasis on differences between dogs and cats in ilarities in the techniques used to manufacture their foods. digestibility of protein, bioavailability of key limiting amino For that reason, petfood processing is addressed in a gener- acids, and control of protein and amino acid metabolism as ic manner and, when appropriate, issues concerning individ- it relates to protein and amino acid requirements and dietary ual species are addressed separately. choice. A new section has been introduced on the use of The information provided in Chapter 13 on food compo- plasma amino acids to assess essential amino acid status of sition, food processing, and other food constituents is the animal. Sufficient data were available to determine that expanded greatly in this volume. Data in Tables 13-1 the protein and amino acid requirements of 4- to 14-week- through 13-8 were compiled from commercial food ingre- old puppies are higher than those puppies over 14 weeks of dient suppliers, published literature, and unpublished data age; so, separate requirements for these different age groups provided by university researchers. The tables include are given. The section on taurine has been updated to means and, when available, the standard deviation and include information on taurine deficiency-induced dilated number (N) of samples used to generate those statistics. cardiomyopathy in the dog and the types of diet that con- Users should examine the standard deviation and N when tribute to this problem. Finally, differences between dogs considering using the mean value as an estimate of the and cats in their responses to amino acid excesses and nutritional content of a specific food ingredient. Obviously, imbalances are discussed. means derived from a large N will better reflect the total Mineral nutrition is covered and presented in terms of 12 population. Means with a large standard deviation may rep- essential macrominerals and trace minerals in Chapter 7. resent the total population but may be a poor estimate for a Acid-base balance is explored for mineral metabolism in specific sample. dogs and cats. Absorption and bioavailability, deficiencies Chapter 14 presents an overview of other food con- and excesses, and nutrient recommendations are discussed stituents that may be used in cat and dog diet formulation. thoroughly for each essential mineral. Additionally, discus- These substances in the diet may affect the structure or func- sion of seven other minerals (arsenic, boron, chromium, tion of the body but their absence from the diet does not fit molybdenum, silicon, nickel, and vanadium) explores their into classical models of nutritional deficiency. Some play potential essentiality, although data on the dog and cat cur- vital roles in the normal metabolism of the animal but can- rently do not clearly establish them as essential. not be deemed essential because the body is capable of syn- The physiologic functions of 14 fat-soluble and water- thesizing them from other dietary components. Other sub- soluble vitamins are described in Chapter 8. This discussion stances that may play a role in the normal function of the includes sections on deficiencies and toxicities for each vita- body must come from an exogenous source, but determina- min. Additional vitamin-like substances, such as inositol, tions of essentiality have yet to be established. Chapter 14 are explored, although none have been demonstrated to be also presents an overview of other food constituents intend- essential for dogs or cats. Finally, vitamin losses in process- ed for technical purposes, such as preservatives. ing and storage of dog and cat foods are examined. Nutrient requirements for growth, adult maintenance, The physiological and biochemical importance of water gestation, and lactation for dogs and cats are provided is outlined in Chapter 9. Regulation of intake, deficiency, in Tables 15-2 through 15-14 (Chapter 15). Estimates are and excess is explored for both dogs and cats. The impor- listed for energy, protein and amino acids, fatty acids, tance of water and feline urinary tract health is discussed. macrominerals, trace minerals, and vitamins.

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4 NUTRIENT REQUIREMENTS OF DOGS AND CATS REFERENCES National Research Council (NRC). 1985. Nutrient Requirements of Dogs. Washington, D.C.: National Academy Press. National Research Council (NRC). 1986. Nutrient Requirements of Cats. Washington, D.C.: National Academy Press. The reader should take note that this document reflects a number of changes made to correct and update an unedited prepublication version of this report. Some val- ues, particularly in Chapter 15, have been revised or deleted based on the availability of new information or to correct errors in calculation. These changes were exam- ined by four independent reviewers and have been approved by the authoring committee and the institu- tion. This final version of the report, therefore, super- sedes the data contained in the prepublication.