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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 11 Nutrient Requirements Table 11-1, which lists estimated minimum nutrient requirements in diets (dry matter basis) for model primate species in six categories, was generated from information presented in the earlier chapters of this volume. Data were sought on model species from eight categories (suborder Strepsirrhini; families Hominidae and Pongidae, Hylobatidae, Cercopithecidae, Cebidae, Callitrichidae, and Tarsiidae; and subfamily Colobinae), but useful data on Hylobatidae and Tarsiidae were not found. The requirement estimates apply to primates fed purified or semipurified diets and assume a high nutrient bioavailability, little to no adverse interaction of nutrients, and an apparent metabolizable energy of 4 kcal·g-1 of dietary dry matter. Energy requirements, as estimated by a variety of techniques, are presented in Chapter 2. As noted by Knapka (2000), many factors influence estimates of nutrient requirements, including genetics, the stage of the life cycle, the rearing environment, the presence of stress, amounts of food consumed, nutrient bioavailability, loss of nutrients between diet formulation and consumption, and criteria of nutritional adequacy. In the studies used to generate Table 11-1, environmental circumstances and the criteria of nutritional adequacy varied greatly. In some instances only one nutrient concentration was tested, or tested nutrient concentrations were very far apart, and that limited the accuracy of the nutrient requirement estimates. Table 11-2 lists estimated dietary nutrient concentrations (dry matter basis) proposed as adequate in diets containing conventional feed ingredients and intended for postweaning primates. The estimated nutrient concentrations in Table 11-2 were based upon primate research reported in previous chapters; nutrient requirements of other herbivorous, omnivorous, and carnivorous mammals published in the National Research Council nutrient requirement series; and the composition of research and commercial primate diets that have successfully supported adult maintenance, reproduction, and growth of young after weaning. The estimates in Table 11-2 are intended to be target levels at the time the diet is fed, and do not account for all potential losses in processing and storage, which can sometimes negatively affect dietary nutrient levels. The nutrient concentrations in Table 11-2 are estimated as adequate but should be used with caution because they may not be appropriate for all species or all postweaning physiologic stages. Nutrient concentrations in Table 11-2 tend to be higher than those in Table 11-1 because the bioavailabilities of nutrients in conventional feed ingredients are usually lower, and nutrient interactions are more likely to require compensation, than when purified or semipurified ingredients are used. Although the bioavailability of all nutrients in natural-ingredient diets should be considered, special attention should be given to phosphorus, zinc, niacin, and biotin. Some of the phosphorus in cereal grains, oilseeds, and their byproducts is found as phytate and is poorly available to simple-stomached animals because of the absence of endogenous digestive enzymes that release phosphorus from its bound form. Phytate also forms complexes with zinc, which render it unavailable for absorption, and additional zinc might be required when phytate is present in the diet. Iron deficiency also has been noted when isolated soy protein replaced casein in semipurified diets fed to rhesus macaques (Macaca mulatta), presumably because of binding by phytate (Fitch et al., 1964). Some natural fiber sources reportedly reduce the activity of amylase, lipase, trypsin, and chymotrypsin in the intestinal tract of human patients with pancreatic insufficiency, although the basis for the inhibition is not entirely clear (Gallaher and Schneeman, 1996). Likewise, some natural sources of fiber reportedly reduce absorption of calcium, magnesium, iron, copper, and zinc; the presence of phytate, rather than fiber itself, might account for some of this reduction (Gallaher and Schneeman, 1996; Jenkins et al., 1999). Some vitamins are bound in organic combinations of limited availability (Ammerman et al., 1995; Jacob and Swendseid, 1996; Mock, 1996). Much of the niacin in cereal
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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 TABLE 11-1 Estimated Nutrient Requirements (in Dietary DM) of Primate Model Species Fed Purified or Semipurified Dietsa Cercopithecidae Cebidae Callitrichidae Colobinae Strepsirrhini Pongidae and Hominidaeb Nutrient Macaque Baboon Squirrel monkey Cebus Howler Marmoset, Tamarin Colobus, Langur Lemur Chimpanzee Humans Crude protein, %c 8m — 8-21g 7m 7-10g 7m 12-18g — — 14gd 6 Taurine, %e — — — — — — — — — — Essential n-3 fatty acids, % f 0.5 — 0.5 0.5 — — — — 0.5 — Essential n-6 fatty acids, % g 2 — 2 2 — — — — 2 — NDF, %h 10 — — — 30 10 30 20 20 — ADF, % i 5 — — — 15 5 15 10 10 — Ca, % 0.55m — — — — — — — — 0.22 P, % 0.33m — — — — — — — — 0.14 Mg, % 0.04m — — — — — — — — 0.074 K, % — 0.24md — — — — — — — — Na, % — 0.25md — — — — — — — — Cl, % — 0.27md — — — — — — — — Fe, mg·kg-1 100g — — — — — — — — 16 Cu, mg·kg-1 15d — — — — — — — — 1.8 Mn, mg·kg-1 44d — — — — — — — — 4.1 Zn, mg·kg-1 20g 13m — 17g — — — — — — 19 I, mg·kg-1 — — — — 0.65d — — — 0.3 Se, mg·kg-1 0.11 — 0.11 — — — — 0.11 Cr+3,mg·kg-1 — — 0.09 — — — — 0.06 Vitamin A, IU·kg-1 5,000 — 12,000d — — — — — — 5,333 Vitamin D3,IU·kg-1 1,000 — 1,250d 1,000 — 2,400d — — — 800 Vitamin E, mg·kg-1 j 68d — — — — >95-130l — — — 30 Vitamin K, mg·kg-1 k >0.06- 3.0 l — — — — — — — — 0.3 Thiamin, mg·kg-1 1.1 — — — — — — — — 2.3 Riboflavin, mg·kg-1 1.7 — — 1.7 — — — — — 2.4 Pantothenic acid, mg·kg-1 20d — 20d — — — — — — 10 Niacin, mg·kg-1 16 — — — — — — — — 30 Vitamin B6,mg·kg-1 4.4d 3.1d — 2-4g — — — — — 2.9 Biotin, mg·kg-1 0.11 — — — — — — — — 0.06 Folacin, mg·kg-1 1.5g — 1.5g 3.3r 1.5g 3.3r — — — — — 0.8 Vitamin B12,mg·kg-1 0.011 0.011 — — — — — — — 0.005 Vitamin C, mg·kg-1 110 — — — — — — — — 170 aEstimated from published data in prior chapters, assuming apparent metabolizable energy at 4.0 kcalg-1 of dry matter, high nutrient bioavailability, and little to no adverse nutrient interactions. Values with following subscripts were derived from studies concerned with maintenance (m) of adults, reproduction (r), or growth (g) of young. Values without a subscript were presumed adequate for all life stages. bFor comparison, recommended dietary allowances or adequate intakes for humans (approximate means of non-reproducing adult age and sex categories), assuming a daily intake of 500 g of dietary dry matter (NRC, 1989 [protein only]; Institute of Medicine, 1997, 1998, 2000, 2001). cProtein requirement depends on amounts and proportions of essential amino acids. Growth requirements decline with age. dLowest concentration tested. eTaurine appears to be required in the diet during the first post-natal year. fEssential n-3 fatty acid requirements met by indicated concentration of a-linolenic acid. If supplied by eicosapentaenoic acid and/or docosahexaenoic acid, required concentration may be less (see Chapter 5). gEssential n-6 fatty acid requirements met by indicated concentration of linoleic acid. hNeutral-detergent fiber. Not a nutrient, but indicated or higher concentration appears to promote gastrointestinal health in indicated primates after weaning (see Chapter 3). iAcid-detergent fiber. Not a nutrient, but indicated or higher concentration appears to promote gastrointestinal health in indicated primates after weaning (see Chapter 3). jAs all-rac-a-tocopheryl acetate. kAs phylloquinone. lLower concentration inadequate, higher concentration adequate. grains and a considerable amount of the niacin in oil seeds is bound and unavailable to simple-stomached animals. Bioavailability of biotin in corn is near 100%, but it is bound and only about 50% available in wheat, barley, triticale, and sorghum grain. Diets comprised of conventional feed ingredients often contain items such as ground grains, grain byproducts, oilseed meals, forage meals, animal byproducts, fats or oils, calcium and phosphorus sources, salt, and vitamin and trace mineral premixes. These or other items are combined
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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 TABLE 11-2 Estimated Adequate Nutrient Concentrations (Dry Matter Basis) in Diets Containing Conventional Feed Ingredients Intended for Post-weaning Nonhuman Primates, Accounting for Potential Differences in Nutrient Bioavailabilities and Adverse Nutrient Interactions, But Not Accounting for Potential Losses in Feed Processing and Storagea Nutrient Concentration Crude protein, % 15-22b Essential n-3 fatty acids, % 0.5 Essential n-6 fatty acids, % 2 NDF, % 10-30c ADF, % 5-15c Ca, % 0.8 Total P, % 0.6d Non-phytate P, % 0.4 Mg, % 0.08 K, % 0.4 Na, % 0.2 Cl, % 0.2 Fe, mg·kg-1 100e Cu, mg·kg-1 20 Mn, mg·kg-1 20 Zn, mg·kg-1 100 I, mg·kg-1 0.35 Se, mg·kg-1 0.3 Trivalent Cr, mg·kg-1 0.2 Vitamin A, IU·kg-1 8,000 Vitamin D3,IU·kg-1 2,500 f Vitamin E, mg·kg-1 100g Vitamin K, mg·kg-1 0.5h Thiamin, mg·kg-1 3.0 Riboflavin, mg·kg-1 4.0 Pantothenic acid, mg·kg-1 12.0 Available niacin, mg·kg-1 25.0i Vitamin B6,mg·kg-1 4.0 Biotin, mg·kg-1 0.2 Folacin, mg·kg-1 4.0 Vitamin B12,mg·kg-1 0.03 Vitamin C, mg·kg-1 200 j Choline, mg·kg-1 750 aBased upon primate research reported in previous chapters; nutrient requirements of other herbivorous, omnivorous, and carnivorous mammals published in the National Research Council nutrient requirement series; and composition of research and commercial primate diets that have successfully supported adult maintenance, reproduction, and growth of young after weaning. These nutrient concentrations have not been directly tested as a group with any primate, and may not be appropriate for all species or all post-weaning physiologic stages. bLactation and growth of young—particularly of smaller primates, such as callitrichids—can be more satisfactory when the higher protein concentrations in this range are used. Required concentrations are greatly affected by protein quality (amounts and proportions of essential amino acids), and this issue must be considered. Taurine appears to be a dietary essential for some primate species through the first postnatal year. cAlthough not nutrients, neutral-detergent fiber (NDF) and acid-detergent fiber (ADF), when used at the concentrations shown in Table 11-1 for the indicated model species, were positively related to gastrointestinal health. dMuch of the phytate phosphorus found in soybean meal and some cereals appears to be of limited bioavailability. eBecause some primates appear to be susceptible to iron-storage disease, particularly in the absence of iron-binding polyphenols found in some plants and when large quantities of fruits are offered, it might be desirable to limit dietary iron concentrations to near or slightly below this concentration. However, this is difficult because of the iron associated with use of calcium phosphates (produced from rock phosphate) as a phosphorus source. Calcium phosphates produced from bone (as a byproduct of gelatin manufacture) are lower in iron. In either case, iron in the phosphate source is thought to be lower in bioavailability than iron in ferrous sulfate, as long as the intake of fruits and their associated citrate and ascorbate contents (which promote iron absorption) is limited. fThere are anecdotal reports of higher vitamin D3 requirements in callitrichids under certain circumstances (see Chapter 7). gAs all-rac- -tocopheryl acetate. hAs phylloquinone. iNiacin in corn, grain sorghum, wheat, and barley is poorly available, as is niacin in byproducts of these grains unless they have undergone fermentation or wet-milling. jAscorbyl-2-polyphosphate is a source of vitamin C that is biologically active and relatively stable during diet extrusion and storage.
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Nutrient Requirements of Nonhuman Primates: Second Revised Edition, 2003 in appropriate formulations, mixed, and are commonly extruded. Sometimes high-moisture fruits and vegetables, browse, or other supplements are fed along with dry extrusions for purposes of environmental enhancement, as described in Chapter 13. Foods used for environmental enhancement should not be used excessively and they should not dilute energy density, which can adversely limit dry matter consumption to below needs. Likewise, these foods should not distort the nutrient balance of a diet formulated to be nutritionally complete. REFERENCES Ammerman, C.B., D.H. Baker, and A.J. Lewis (Eds.). 1995. Bioavailability of Nutrients for Animals. San Diego: Academic Press. 441 pp. Fitch, C.D., W.E. Harville, J.S. Dinning, and F.S. Porter. 1964. Iron deficiency in monkeys fed diets containing soybean protein. Proc. Soc. Exp. Biol. Med. 116:130-133. Gallaher, D.D., and B.O. Schneeman. 1996. Dietary fiber. Pp. 87-97 in Present Knowledge in Nutrition, 7th ed., E.E. Ziegler and L.J. Filer, Jr., Eds. Washington, DC: ILSI Press. Institute of Medicine. 1997. Dietary Reference Intakes for Calcium, Phosphorus, Magnesium, Vitamin D, and Fluoride. Washington, DC: National Academy Press. Institute of Medicine. 1998. Dietary Reference Intakes for Thiamin, Riboflavin, Niacin, Vitamin B6, Folate, Vitamin B12, Pantothenic Acid, Biotin, and Choline. Washington, DC: National Academy Press. Institute of Medicine. 2000. Dietary Reference Intakes for Vitamin C, Vitamin E, Selenium, and Carotenoids. Washington, DC: National Academy Press. Institute of Medicine. 2001. Dietary Reference Intakes for Vitamin A, Vitamin K, Arsenic, Boron, Chromium, Copper, Iodine, Iron, Manganese, Molybdenum, Nickel, Silicon, Vanadium, and Zinc. Washington, DC: National Academy Press. Jacob, R.A., and M.E. Swendseid. 1996. Niacin. Pp. 184-190 in Modern Nutrition in Health and Disease, 9th ed., M.E. Shils, J.A. Olson, M. Shike, and A.C. Ross, Eds. Baltimore: Williams & Wilkins. Jenkins, D.J.A., T.M.S. Wolever, and A.L. Jenkins. 1999. Fiber and other dietary factors affecting nutrient absorption and metabolism. Pp. 679-698 in Modern Nutrition in Health and Disease, 9th ed., M.E. Shils, J.A. Olson, M. Shike, and A.C. Ross, Eds. Baltimore: Williams & Wilkins. Knapka, J.J. 2000. Factors influencing required dietary nutrient concentrations. Lab. Anim. 29:47-50. Mock, D.M. 1996. Biotin. Pp. 220-235 in Present Knowledge in Nutrition, 7th ed., E.E. Ziegler and L.J. Filer, Jr., Eds. Washington, DC: ILSI Press. National Research Council. 1989. Recommended Dietary Allowances, 10th ed. Washington, DC: National Academy Press.
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