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Nutrient Requirements (J of the Young Calf From birth until weaning to dry feed, the calf undergoes tremendous physiologic and metabolic changes (Toullec and Guilloteau, 19891. During the preruminant stage, digestion and metabolism are similar to those of nonrumi- nant animals in many respects. Thus, dietary requirements are best met with high-quality liquid diets formulated from sources of carbohydrates, proteins, and fats that are digested efficiently. The most critical period is the first 2-3 wk of life, during which time the calms digestive system is immature but developing rapidly with regard to digestive secretions and enzymatic activity (Toullec and Guilloteau, 1989; Davis and Drackley, 19981. Calves raised for purposes other than veal production should be encouraged to consume dry feed at an early age to stimulate development of a functional rumen. Develop- ment of the ruminal epithelial tissue that is responsible for absorption of volatile fatty acids (VFA) depends on the presence of the VFA, particularly butyrate (Sander et al., 19591. The chemical composition and physical form of the starter feed are important characteristics (Warner, 19911. The starter should be relatively high in readily fermentable carbohydrates but adequate in digestible fiber to support the fermentation necessary for proper ruminal tissue growth (Brownlee, 1956; Flatt et al., 1958; Williams and Frost, 1992; Greenwood et al., 19971. The rumen and its microbial population are immature at this stage (Anderson et al., 1987a,b) and ruminal cellulose digestibility is limited (Williams and Frost, 19921. Consequently, long hay is not as effective as concentrates in developing a functional rumen and limits metabolizable energy intake in young calves (Stobo et al., 19661. Long hay should not be fed to calves until after weaning (Quigley, 1996a; Davis and Drackley, 19981. Nevertheless, adequate particle size of starter feed whether pelleted, ground, ortexturized is important to prevent abnormal development and keratini- zation of ruminal papillae and to prevent impaction of fine particles between papillae (McGavin and Morrill, 1976; Greenwood et al., 1997; Beharka et al., 19981. With respect to the nutrient requirements of the calf, three phases of development related to digestive function are recognized (Davis and Clark, 19811: Liquid-feeding phase. All or essentially all the nutrient requirements are met by milk or milk replacer. The quality of these feeds is preserved by a functional esophageal groove, which shunts liquid feeds directly to the abomasum and so avoids microbial breakdown in the reticulo-rumen (Orskov, 19721. Transition phase. Liquid diet and starter both contrib- ute to meeting the nutrient requirements of the calf. Ruminant phase. The calf derives its nutrients from solid feeds, primarily through microbial fermentation in the reticulo-rumen. This chapter discusses nutrient requirements of calves in each of those phases. ENERGY REQUIREMENTS OF CALVES Energy requirements of calves, like those of other ages and classes of cattle, can be expressed in numerous ways (see Chapter 21. Regardless of the system preferred, it is imperative to understand where the major losses of energy occur as the energy-yielding components of the diet undergo digestion and metabolism. If the efficiencies of conversion of gross energy to digestible energy or metabo- lizable energy and of conversion of metabolizable energy to net energy (both NEM and NEG) are known, users can select the system that best fits their needs. In this edition, the energy requirements of calves have been derived on the basis of metabolizable energy; how- ever, requirements and feed composition also are given in units of net energy and digestible energy for those who prefer to use those systems. Data on energy requirements are organized around replacement calves fed only milk or milk replacer (Table 10-11; calves fed milk and starter feed or milk replacer and 214

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Nutrient Requirements of the Young Calf 215 TABLE 10-1 Replacer Daily Energy and Protein Requirements of Young Replacement Calves Fed Only Milk or Milk Energy Protein Live Weight Gain Dry Matter NEMb NEGC MEd DEe ADPf CPg Vitamin Ah (kg) (g) Intakea (kg) (Meal) (McCoy) (McCoy) (Meal) (g) (g) (IU) 25 0 0.24 0.96 0 1.12 1.17 18 20 2,750 200 0.32 0.96 0.26 1.50 1.56 65 70 2,750 400 0.42 0.96 0.60 2.00 2.08 113 121 2,750 30 0 0.27 1.10 0 1.28 1.34 21 23 3,300 200 0.36 1.10 0.28 1.69 1.76 68 73 3,300 400 0.47 1.10 0.65 2.22 2.31 115 124 3,300 40 0 0.34 1.37 0 1.59 1.66 26 28 4,400 200 0.43 1.37 0.31 2.04 2.13 73 79 4,400 400 0.55 1.37 0.72 2.63 2.74 120 129 4,400 600 0.69 1.37 1.16 3.28 3.41 168 180 4,400 45 0 0.37 1.49 0 1.74 1.81 28 30 4,950 200 0.46 1.49 0.32 2.21 2.30 76 81 4,950 400 0.59 1.49 0.75 2.82 2.94 123 132 4,950 600 0.74 1.49 1.21 3.50 3.64 170 183 4,950 50 0 0.40 1.62 0 1.88 1.96 31 33 5,500 200 0.45 1.62 0.34 2.37 2.47 78 84 5,500 400 0.63 1.62 0.77 3.00 3.13 125 135 5,500 600 0.78 1.62 1.26 3.70 3.86 173 185 5,500 eddy matter intake necessary to meet ME requirements for calves fed milk replacer composed primarily of milk proteins and containing ME at 4.75 Mcal/kg of day matter. bNEM (Meal) = 0.086 LW075, where LW is live weight in kilograms. ONES (Meal) = (0.84 LW0355 X LWGi2) X 0.69, where LW and LWG (live weight gain) are in kilograms. l aME (Meal) = 0.1 LW075 + (0.84 LW0355 X LWGi2), where LW and LWG are in kilograms. eDE (Meal) = ME/0.96. fADP(apparent digestible prote~n,gid) = 6.25[1/BV(E + G + M X D)M X D].BV(biologiCvalue)isassumedtobeO.8.E(endogenousunnalynitrogen)isO.2LW75/d, where LW is in kilograms. M (metabolic fecal nitrogen) is 1.9 g/kg of dry matter intake (D). G (nitrogen in live weight gain) is 30 g/kg of LWG. gCP (crude protein) = ADP/0.93. The digestibility of undenatured milk proteins is assumed to be 93 percent. hVitamin A (IU) = 110 IU/kg of LW. See Chapter 7. starter feed (Table 10-21; calves reared for veal on only milk or milk replacers (Table 10-31; and weaned replace- ment calves to 100 kg of body weight fed starter or grower diets (Table 10-41. The amount of liquid feed (milk or milk replacer) offered to replacement calves is restricted to encourage intake of dry feed (starter), but calves reared for veal are fed milk or milk replacer at near ad libitum intakes. Young Replacement Calves Fed Milk or Milk Replacer Only The energy requirements of young calves fed only milk or milk replacer and weighing 25-50 kg are given in Table 10-1. On the basis of available data, NEM is set at 0.086 Mcal/kg075 of live weight (LOO) daily as in the previous edition of this publication (National Research Council, 19891. This equates reasonably well with estimates of fast- ing metabolism of young milk-fed calves that are limited in activity (see chapter 4 of Davis and Drackley, 19981. The efficiency of use of metabolizable energy (ME) from milk or milk replacer to meet maintenance requirements is set at 86 percent. Consequently, maintenance ME is defined as 0.100 Mcal/kg075 daily. The values for ME and efficiency of ME use for maintenance are within the range of values in the scientific literature (Van Es et al., 1969; Johnson and Elliott, 1972a,b; Holmes and Davey, 1976; Okamoto et al., 1986; Arieli et al., 1995; Gerrits et al., 19961. The Agricultural Research Council (ARC) specified an ME requirement of 0.102 Mcal/kg075 daily, with an efficiency of use of ME for maintenance of 85 percent (Agricultural Research Council, 19801. Requirements for ME were calculated with the equation derived by Toullec (1989) as follows: ME requirement (Meal/d) = 0.1 LW075 + (0.84 LW0355~LWG~.2> (10-1) where LW and daily liveweight gain (LWG) are in kilo- grams. The first portion of the equation sets the ME required for maintenance at 100 kcal/kg075 per day. The second portion of the equation is used to derive the ME required for LWG, which is a function of both body size (LOO) and rate of gain (LWG). This equation was derived on the basis of an efficiency of conversion of ME to NEG of 69 percent for calves fed only milk or milk replacer, which is consistent with most published values (Gonzalez- ;Jimenez and Blaxter, 1962; Van Es et al., 1969; Johnson and Elliott, 1972a,b; Vermorel et al., 1974; Webster et al., 1975; Donnelly and Hutton, 1976a,b; Holmes and Davey, 1976; Neergard, 1976; Toullec, 1989; Gerrits et al., 19961. The energy content of LWG predicted by equation 10-1 is 1556 kcal/kg LWG for a 40-kg calf gaining 200 g/d, and 2567 kc al/kg LWG for a 75-kg calf gaining 800 g/d. Values

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Live Weight Gain (kg) (g) 216 Nutrient Requirements of Dairy Cattle TABLE 10-2 Daily Energy and Protein Requirements of Calves Fed Milk and Starter or Milk Replacer and Starter Energy Dry Matter Intakea (kg) NEMb (Meal) NEGC (Meal) o 0.28 0.65 o 0.30 0.68 o 0.31 0.72 1.16 o 0.32 0.75 1.21 o 0.34 0.77 1.26 1.78 o 0.35 0.80 1.30 1.84 o 0.36 0.83 1.34 1.90 Protein MEd (Meal) 1.34 1.77 2.33 1.50 1.96 2.55 1.66 2.14 2.76 3.44 1.81 2.31 2.96 3.67 1.96 2.48 3.15 3.89 4.69 2.11 2.64 3.33 4.10 4.93 2.25 2.80 3.51 4.31 5.16 DEe (Meal) ADPf (g) CPg (g) 26 84 141 29 87 145 33 90 148 205 36 93 151 209 38 96 154 212 270 41 99 157 215 273 44 102 159 217 275 Vitamin Ah (IU) 30 35 40 45 50 55 60 o 200 400 200 400 200 400 600 200 400 600 o 200 400 600 800 200 400 600 800 o 200 400 600 800 0.32 0.42 0.56 0.36 0.47 0.61 0.40 0.51 0.66 0.83 0.44 0.56 0.71 0.88 0.47 0.60 0.76 0.94 1.13 0.51 0.63 0.80 0.99 1.18 0.54 0.67 0.84 1.04 1.24 1.10 1.10 1.10 1.24 1.24 1.24 1.37 1.37 1.37 1.37 1.49 1.49 1.49 1.49 1.62 1.62 1.62 1.62 1.62 1.74 1.74 1.74 1.74 1.74 1.85 1.85 1.85 1.85 1.85 1.43 1.89 2.49 1.61 2.09 2.73 1.78 2.29 2.95 3.68 1.94 2.47 3.16 3.93 2.10 2.65 3.37 4.17 5.02 2.25 2.83 3.57 4.39 5.27 2.41 3.00 3.76 4.61 5.52 23 72 122 25 75 125 25 78 128 178 31 80 130 180 33 83 133 183 233 36 85 135 185 236 38 88 138 188 238 3,300 3,300 3,300 3,850 3,850 3,850 4,400 4,400 4,400 4,400 4,950 4,950 4,950 4,950 5,500 5,500 5,500 5,500 5,500 6,O50 6,O50 6,O50 6,O50 6,O50 6,600 6,600 6,600 6,600 6,600 aThese data apply to calves fed milk replacer (MR) plus starter. MR contains ME at 4.75 Mcal/kg of DM and starter ME at 3.28 Mcal/kg. It is assumed that MR provided 60 percent and starter 40 percent of dry matter intake; thus, dry matter consumed contained ME at 4.16 Mcal/kg. The DMI here is the total necessary to meet ME requirements and is not intended to predict voluntary intake. NEM (Meal) = 0.086 LW075, where LW is live weight in kilograms. ONES (Meal) = (0.84 LW0355 X LWGi2) X 0.69, where LW and LW gain (LWG) are in kilograms. TIME (Meal) was computed as follows: ME (maintenance) = NEM/0.825. Efficiency of use of ME for maintenance (0.825) was computed as average of efficiencies of 0.86 for MR and 0.75 for starter, weighted according to proportions of ME supplied by each feed. ME (gain) = NEJO.652. Efficiency of use of ME for gain (0.652) was computed as weighted average of efficiencies of 0.69 and 0.57 for MR and starter, respectively. eDE (Meal) = ME/0.934. Efficiency of conversion of DE to ME is assumed to be 0.96 for MR and 0.88 for starter. fADP (apparent digestible protein, g/d) = 6.25 [1/BV(E + G + M X D)- M X D]. BV (biologic value) = 0.764 (weighted average of MR = 0.8 and starter = 0.70); E (endogenous urinary nitrogen, g ) = 0.2LW75; G (nitrogen content of gain, g) = 30 g/kg gain; M (metabolic fecal nitrogen, g/d) = 2.46 X dry matter intake, D, kg). Metabolic fecal nitrogen for MR assumed to be 1.9 g/kg of DMI and for starter 3.3 g/kg of DMI. gCP (crude protein, g) = ADP/0.8645. Digestibility of protein was assumed to be weighted average of 93 percent for MR and 75 percent for starter; MR was assumed to contain 21 percent CP and starter 18 percent CP. hVitamin A (IU) = 110 IU/kg of LW. See Chapter 7. predicted by this equation are similar to those in the 1989 edition of this publication for smaller calves at low rates of gain (1460 kc al/kg LWG for a 40-kg calf gaining 200 g/ d) but are substantially higher than the 1989 edition for larger calves at higher rates of gain (1869 kcal/kg LWG for a 75-kg calf gaining 800 g/d). Values predicted by the present equation agree well with available experimental data on body composition of dairy calves (Webster et al., 1975; Donnelly and Hutton, 1976b; Holmes and Davey, 1976; Neergard, 1976; Gerrits et al., 19961. Data for com- position of LWG for dairy calves of current genotypes would be useful for future refinement of requirements for growth. The ME requirements given in Table 10-1 for calves weighing 30-60 kg and gaining at different rates are in close agreement with most published data. The digestible energy (DE) values in Table 10-1 are calculated from ME, assuming an efficiency of 96 percent for conversion of DE to ME (Neergard, 1976; National Research Council, 1989; Toullec, 1989; Gerrits et al., 19961. Users that desire requirements for higher rates of gain than included in Table 10-1 for calves fed milk or milk replacer only should refer to Table 10-3. Users should be aware that ME requirements for main- tenance may be underestimated for calves during the first week of life because of the high and variable basal meta-

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Nutrient Requirements of the Young Calf 217 TABLE 10-3 Daily Energy and Protein Requirements of Veal Calves Fed Only Milk or Milk Replacer Energy Protein Live Gain Dry Matter NEMb NEGC MEd DEe ADPf CPg Vitamin Ah Weight (kg) (g) Intakea (kg) (Meal) (Meal) (Meal) (Meal) (g) (g) (IU) 40 0 0.34 1.37 0 1.59 1.66 26 28 4,400 300 0.49 1.37 0.51 2.32 2.42 97 104 4,400 600 0.69 1.37 1.16 3.28 3.41 168 180 4,400 50 0 0.40 1.62 0 1.88 1.96 31 33 5,500 300 0.56 1.62 0.55 2.67 2.79 102 109 5,500 600 0.78 1.62 1.26 3.71 3.86 172 185 5,500 900 1.02 1.62 2.05 4.85 5.05 244 262 5,500 60 0 0.45 1.85 0 2.16 2.25 35 38 6,600 300 0.63 1.85 0.58 3.00 3.13 106 114 6,600 600 0.86 1.85 1.34 4.10 4.27 177 190 6,600 900 1.12 1.85 2.18 5.32 5.54 248 267 6,600 70 0 0.51 2.08 0 2.42 2.52 39 42 7,700 300 0.70 2.08 0.62 3.32 3.45 110 119 7,700 600 1.94 2.08 1.42 4.48 4.66 181 195 7,700 900 1.21 2.08 2.31 5.76 6.01 253 272 7,700 1,200 1.50 2.08 3.26 7.14 7.44 324 348 7,700 80 0 0.56 2.30 0 2.68 2.79 44 47 8,800 300 0.76 2.30 0.65 3.61 3.76 115 123 8,800 600 1.02 2.30 1.49 4.83 5.03 186 200 8,800 900 1.30 2.30 2.42 6.18 6.44 257 276 8,800 1,200 1.61 2.30 3.42 7.63 7.95 328 353 8,800 90 0 0.62 2.51 0 2.92 3.04 48 51 9,900 300 0.82 2.51 0.68 3.90 4.06 119 128 9,900 600 1.09 2.51 1.55 5.17 5.39 190 204 9,900 900 1.38 2.51 2.55 6.62 6.85 263 283 9,900 1,200 1.70 2.51 3.56 8.09 8.42 332 357 9,900 100 0 0.67 2.72 0 3.16 3.29 52 55 11,000 300 0.88 2.72 0.70 4.18 4.35 122 132 11,000 600 1.16 2.72 1.61 5.50 5.72 194 208 11,000 900 1.46 2.72 2.62 6.96 7.25 265 285 11,000 1,200 1.80 2.72 3.70 8.52 8.88 336 362 11,000 1,500 2.14 2.72 4.84 10.17 10.59 408 438 11,000 110 0 0.72 2.92 0 3.40 3.54 55 60 12,100 300 0.94 2.92 0.72 4.45 4.63 126 136 12,100 600 1.22 2.92 1.66 5.81 6.05 198 212 12,100 900 1.54 2.92 2.71 7.32 7.63 269 289 12,100 1,200 1.88 2.92 3.83 8.94 9.32 340 366 12,100 1,500 2.24 2.92 5.00 10.65 11.09 412 443 12,100 120 0 0.76 3.12 0 3.63 3.78 59 64 13,200 300 0.99 3.12 0.75 4.71 4.91 130 140 13,200 600 1.29 3.12 1.72 6.12 6.39 201 217 13,200 900 1.62 3.12 2.80 7.68 8.00 273 293 13,200 1,200 1.97 3.12 3.69 9.34 9.74 329 353 13,200 1,500 2.34 3.12 5.16 11.10 11.56 416 447 13,200 130 0 0.81 3.31 0 3.85 4.01 63 67 14,300 300 1.05 3.31 0.77 4.97 5.17 134 144 14,300 600 1.35 3.31 1.77 6.41 6.68 205 220 14,300 900 1.69 3.31 2.88 8.02 8.35 276 297 14,300 1,200 2.05 3.31 4.06 9.74 10.14 348 374 14,300 1,500 2.43 3.31 5.31 11.54 12.02 420 451 14,300 140 0 0.86 3.50 0 4.07 4.24 66 71 15,400 300 1.10 3.50 0.79 5.22 5.43 137 148 15,400 600 1.41 3.50 1.82 6.70 6.98 209 224 15,400 900 1.76 3.50 2.95 8.35 8.70 280 301 15,400 1,200 2.13 3.50 4.17 10.11 10.53 352 378 15,400 1,500 2.52 3.50 5.45 11.97 12.45 423 455 15,400 150 0 0.90 3.69 0 4.29 4.46 70 75 16,500 300 1.15 3.69 0.81 5.46 5.69 141 152 16,500 600 1.47 3.69 1.86 6.98 7.27 212 228 16,500 900 1.82 3.69 3.02 8.67 9.03 284 305 16,500 1,200 2.21 3.69 4.27 10.48 10.91 355 382 16,500 1,500 2.61 3.69 5.58 12.38 12.90 427 459 16,500 aThe DMI necessary to meet ME requirements when veal calves are fed a milk replacer containing ME at 4.75 Mcal/kg of DM. bNEM (Meal) = 0.086 LW075, where LW is live weight in kilograms. CNEG (Meal) = (0.84 LW0355 X LWGi2) X 0.69, where LW and LW gain (LWG) are in kilograms. IMP (Meal) = ().1 LAW' + (0.84 LW0355 X LWGi2), where LW and LWG are in kilograms. eDE (Meal) = ME/0.93. fADP (apparent digestible protein, g/d) = 6.25 [1/BV(E + G + M X D)M X D]. BV (biologic value) is assumed to be 0.8. E (endogenous urinary nitrogen) is 0.2 LW075/d, where LW is in kilograms. M (metabolic fecal nitrogen) is 1.9 g/kg of dry matter intake (D). G (nitrogen in live weight gain) is 30 g/kg LWG. gCP (crude protein) = ADP/0.93. The digestibility of undenatured milk proteins is assumed to be 93 percent. hVitamin A (IU) = 110 IU/kg of LW. See Chapter 7. '7 _ . . . ~ - -

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218 Nutrient Requirements of Dairy CattIe TABLE 10-4 Daily Energy and Protein Requirements of Weaned (Ruminant) Calvesa Energy Protein Live Weight Gain Day Matter NEMb NEGC MEd DEe ADpf CPg Vitamin Ah (kg) (g) Intake (kg) (Meal) (McCoy) (Meal) (McCoy) (g) (g) (IU) 50 0 0.70 1.62 0 2.16 2.58 40 53 5,500 400 1.13 1.62 0.77 3.51 3.92 151 201 5,500 500 1.27 1.62 1.01 3.93 4.35 179 238 5,500 600 1.86 1.62 1.26 4.36 4.77 207 276 5,500 60 0 0.80 1.85 0 2.47 2.89 46 61 6,600 400 1.26 1.85 0.83 3.92 4.33 156 209 6,600 500 1.41 1.85 1.08 4.36 4.77 185 246 6,600 600 1.56 1.85 1.34 4.83 5.23 213 284 6,600 700 1.71 1.85 1.62 5.31 5.70 241 322 6,600 800 1.87 1.85 1.90 5.80 6.19 269 359 6,600 70 0 0.90 2.08 0 2.77 3.19 51 68 7,700 400 1.39 2.08 0.87 4.31 4.71 163 217 7,700 500 1.54 2.08 1.14 4.77 5.17 191 254 7,700 600 1.70 2.08 1.42 5.26 5.66 219 292 7,700 700 1.86 2.08 1.71 5.77 6.16 247 330 7,700 800 2.03 2.08 2.00 6.29 6.67 275 367 7,700 80 0 0.99 2.30 0 3.07 3.48 57 75 8,800 400 1.51 2.30 0.92 4.67 5.07 168 224 8,800 500 1.66 2.30 1.20 5.16 5.56 196 262 8,800 600 1.83 2.30 1.49 5.68 6.07 225 300 8,800 700 2.00 2.30 1.79 6.21 6.59 253 337 8,800 800 2.18 2.30 2.10 6.75 7.13 281 375 8,800 90 0 1.16 2.51 0 3.35 3.76 62 82 9,900 600 2.09 2.51 1.55 6.07 6.46 231 309 9,900 700 2.28 2.51 1.87 6.62 7.00 260 346 9,900 800 2.48 2.51 2.19 7.19 7.57 288 385 9,900 900 2.68 2.51 2.52 7.78 8.15 317 423 9,900 100 0 1.25 2.72 0 3.63 4.04 68 90 11,000 600 2.22 2.72 1.61 6.45 6.83 237 316 11,000 700 2.42 2.72 1.94 7.02 7.40 265 354 11,000 800 2.63 2.72 2.27 7.62 7.99 294 392 11,000 900 2.84 2.72 2.62 8.22 8.59 323 430 11,000 aThese data apply to small-breed female calves from 50 to 80 kg gaining 0.4 to 0.5 kg/d arid large-breed calves from 60 to 100 kg gaining from 0.6 to 0.9 kg/d. bNEM (Meal) = 0.086 LW075 (NRC 1989), where LW is live weight in kilograms. ONES (Meal) = (0.84 LW0355 X LWGi2) X 0.69, where LW and LW gain (LWG) are in kilograms. ~ME, maintenance (Meal) = NEM/0.75. ME values of diets (Meal/kg of DM) are 3.10 for calves weighing 60, 70, and 80 kg and 2.90 for calves weighing 90 and 100 kg. ME, gain (Meal) = NEG/0.57. Sum of ME values for maintenance Flus vain equals total ME requirement. ~ O ~ eDE (Meal) = (ME + 0.45) /1.01 (see Chapter 2). JADP (apparent digestible protein, g/d) as follows: ADP (g/d) = 6.25 [1/BV(E + G + M X D)M X D] where BV is biologic value set at 0.70; E (endogenous urinary nitrogen) = 0.2LW75; G is nitrogen content of gain, assuming 30 g/kg of gain; and M is metabolic fecal nitrogen computed as 3.3 g/kg of dry matter consumed (D). gCP (crude protein) calculated as ADP/0.75. hVitamin A (IU) = 110 IU/kg of LW. See Chapter 7. boric rate observed during this time (Roy et al., 1957; Vermorel et al., 1983; Okamoto et al., 1986; Schrama et al., 1992; Ortigues et al., 1994; Arieli et al., 19951. Further- more, because the digestive tract is immature and develop- ing rapidly, the metabolizability of diets may be lower during this time (Schema et al., 1992; Arieli et al, 1995), thereby overestimating dietary energy supply. The net result of these effects is that LWG of calves during the first week of life may be considerably less than the pre- dicted energy-allowable gains shown in Table 10-1. As more data become available it may become possible in future editions to model these effects. Energy requirement values for young calves in this edi- tion represent several improvements over the previous edi- tion (National Research Council, 19891. First, tabulated values in this edition are derived directly from the equa- tions presented, in contrast with values given in the tables of the 1989 edition that could not be calculated from the information provided. Second, as discussed above, values for the energy content of body weight gain (NEG) in this edition agree more closely with available data on calves derived from slaughter experiments; values in the 1989 edition were too low (see Davis and Drackley, 19981. Third, the equations used to derive the NEM and NEG values for milk or milk replacers in the previous edition were those of Garrett (see National Research Council, 1989) estab- lished for feedlot cattle fed diets with ME content of 2.19- 2.86 Mcal/kg of dry matter (DM). Those equations result in erroneously low NE values for diets of milk or milk- derived products. Garrett (1980) cautioned against using

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Nutrient Requirements of the Young Calf 219 the established equations to derive NE values for foodstuffs with ME values outside the range stated above. A different approach has been taken in this edition to derive the NE values for liquid diets and starter. Young Replacement Calves Fed Milk and Starter Feed or Milk Replacer and Starter Feed Under good management on dairy farms, calves should be consuming appreciable nutrients from starter feed by the second week of life. To encourage early consumption of calf starter, calves should be given free access to water and a nutritious, highly palatable starter feed from the first week of life until they are weaned. Consumption of starter feed is critical to development of an active, functioning rumen. Fermentation products, principally butyrate, from fermentation of solid feeds in the developing rumen are responsible for development of functional ruminal epithe- lial tissue (Sander et al., 19591. Deriving the energy requirements of calves fed a combi- nation diet (liquid plus dry feed) requires the application of basic knowledge from related areas because there are few data on the subject. Only one study, which used three calves per treatment, has examined this question directly by using calorimetry (Holmes and Davey, 19761. The main- tenance requirement and efficiency of use of ME by calves did not differ appreciably between an all-milk diet and a diet consisting of milk and dry feed. Regardless of the diet fed, the NE required for mainte- nance and gain should not change. Efficiencies of utiliza- tion of ME for maintenance and gain will be somewhat lower for starter feeds than for milk or milk replacer (National Research Council, 19781. As described for Table 10-1, calves use the ME from milk or milk replacer with efficiencies of 86 percent and 69 percent for maintenance and gain, respectively. Efficiency of ME use from milk or milk replacer is assumed not to change when starter also is consumed. The previous edition of this publication (National Research Council, 1989) used the equations of Garrett (1980) to derive the efficiencies of utilization of ME (percent) from starter for maintenance (km) and gain (kg): km = 51.045 ME 10.836 ME2 + 0.754 ME3 - 7.35 (10-2) kg = 76.149 ME 15.755 ME2 + 1.062 ME3 - 69.7 (10-3) where ME is expressed as Mcal/kg DM. However, these data were for older growing cattle fed feedlot diets and are not appropriate for young calves. For example, the Garrett (1980) equations yield efficiencies of ME use for maintenance and gain of 69.4 and 46.4 percent, respectively, for a starter containing 3.1 Mcal ME/kg DM. These efficiencies are lower than those calculated from experimental data (Holmes and Davey, 1976) and used in other systems (Agricultural Research Council, 19801. Furthermore, the Garrett (1980) equations were developed using ME values calculated as 0.82 DE (National Research Council, 19891. Because methane production is minimal even in young calves consuming 44 percent of their ME from concentrates (Holmes and Davey, 1976), these derived ME values are too low when compared with experi- mental data (Spanski et al., 19971. Consequently, the use of the Garrett (1980) equations for young calves has been discontinued in this edition. The Agricultural Research Council (1980) calculated efficiencies of ME use for maintenance and gain as a func- tion of the metabolizability (ME/GE, or "q") of the diet. Over the range of ME concentrations expected for calf starters and growers (2.5-3.4 Mcal/kg), the efficiency of ME use for maintenance would vary from only about 72 to 77 percent, and that for gain from 50 to 59 percent. In this edition, efficiencies of ME use from dry feeds for maintenance and gain were fixed at 75 and 57 percent, respectively. The efficiency of use of ME from the total diet is then calculated as the average of individual eff~cienc- ies for milk and starter, weighted according to their contri- bution to the total ME in the diet. In the example given in Table 10-2, it was assumed that a calf at about 2 wk of age would consume on the average a diet in which 60 percent of DM intake (DMI) is derived from milk replacer (ME at 4.75 Me al/kg of DM) and 40 percent from starter (ME at 3.28 Mcal/kg of DM). In this diet, milk-replacer supplies 68 percent of the total ME, and starter supplies 32 percent. Consequently, the overall efficiencies for use of ME in the combined diet (milk replacer plus starter) are 82.5 and 65.2 percent for mainte- nance and gain, respectively, calculated as the weighted average (weighted by contribution to the total ME supply) of the individual efficiencies. The computer model included with this edition calculates these values for varied proportions of DMI from milk and starter or milk replacer and starter. A comparison of the ME requirement of a 50-kg calf gaining 400 g/d when fed only milk or milk replacer (see Table 10-1) with the ME requirement of the same calf fed milk and starter or milk replacer and starter (Table 10-2) reveals a relatively small difference (3.00 vs 3.15 Mcal/d). The ME requirements given here for calves consuming both starter and milk or milk replacer are markedly lower than those given in the 1989 edition (5.90 Mcal/d) but are similar to those given by Roy (19801. A comparison of LWG predicted by this model with actual performance of calves receiving both milk or milk replacer and starter in 16 published research studies reveals good agreement (Stewart and Schingoethe, 1984; Jenny et al., 1991; laster et al., 1992; Reddy et al., 1993; Akayezu et al., 1994; Quigley et al.. 1994a Abdelgadir and Morrill, 1995; Quigley et al.,

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220 Nutrient Requirements of Dairy Cattle 1995; Abdelgadir et al., 1996a, b; Quigley, 1996b; Quigley and Bernard, 1996; Quigley and Welborn, 1996; Terui et al., 1996; Quigley et al., 1997b; Lammers et al., 19981. Table 10-2 also presents requirements for energy in units of DE. Values for DE were calculated as ME/0.934, representing the weighted average of conversion of DE to ME for milk or milk replacer (0.96) and starter (0.881. The conversion from ME to DE for starter was calculated as (ME + 0.451/1.01 (National Research Council, 1989), as described in Chapter 2 (also see later discussion on energy values for feeds). The DMI listed in Tables 10-1 through 10-4 have been computed as the amount of DM necessary to provide the ME requirement. Consequently, these should not be con- strued to be predictions of voluntary feed intake. An analy- sis of literature data presented elsewhere (see chapter 16 of Davis and Drackley, 1998) predicts that intake of DM from starter increases from about 0.8-1.0 percent of BW at 3 wk of age to about 2.8-3.0 percent of BW at 8 wk of age. Veal Calves The calculations used to derive the ME requirements for veal calves (Table 10-3) are the same as those for milk- fed replacement calves (Table 10-11. Veal calves are fed essentially for ad libitum intake, so rates of gain will be higher than those of limit-fed replacement calves. The ME and DM requirements given here agree closely with those reported by Webster et al. (1975) on the basis of an energy- balance study with veal calves. Ruminant Calves (Large-Breed and Small-Breed FemalesJ from Weaning to Body Weight of 100 Kilograms In the previous edition of Nutrient Requirements of Dairy Cattle, (National Research Council, 1989) no infor- mation was given on the nutrient requirements of calves from weaning to 100 kg of body weight even though this is a critical period in the life ofthe replacement calf. Similar to calves consuming milk and starter, very few research data determined by calorimetry or comparative slaughter studies exist for this class of cattle. However, the subcom- mittee believes that estimates should be made. Methods used in this edition to establish requirements for growth of heifers from 100 to 500 kg of body weight could not be extrapolated accurately to calves weighing less than 100 kg. Given the paucity of data on tissue growth and nutrient use for this class of calves, estimated requirements have been derived using the same methodology as described already for younger calves. Users will note that require- ments for ruminant calves weighing less than 100 kg do not merge smoothly into requirements for larger calves. Energy-allowable LWG was predicted using this model from LW and estimated ME intakes from 25 treatments in 19 published studies (Stewart and Schingoethe, 1984; Beharka et al., 1991; Chester-;[ones et al., 1991; Jenny et al., 1991; Quigley et al., 1991; Quigley et al., 1992; Reddy et al., 1993; Akayezu et al., 1994; Jackson and HemLen, 1994; Kuehn et al., 1994; Maiga et al., 1994; Quigley et al., 1994a; Abdelgadir and Morrill, 1995; Abdelgadir et al., 1996a,b; Quigley, 1996b; Terui et al., 1996; Kincaid et al., 19971. Comparisons were expressed as predicted/observed; the mean was 1.04. Twelve predicted values were greater than observed, twelve were less than observed, and one was equal. As more research information becomes available, future editions of this publication may be better able to define requirements for this group of calves. However, in comparing requirements established here with literature data on average daily gains, the methodology presented in this edition adequately predicts gains of large-breed calves up to 100 kg and small-breed calves to 80 kg. Table 10-4 shows the requirements of weaned calves weighing 50-100 kg and gaining at various rates. Calves weighing 50-80 kg were assumed to be fed a starter con- taining ME at 3.1 Meal of ME per kg of DM, and those weighing 90-100 kg a starter or grower containing ME at 3.0 Meal per kg of DM. Given the paucity of data, no distinction is made between large and small breed calves. Similarly, no distinction is made between male and female calves since differences are negligible before about 100 kg LW (National Research Council, 19781. Elects of Environmental Temperature on Energy Requirements of Young Calves The calf is born with limited body energy reserves and only modest insulation afforded by hair coat and body fat. A newborn calf is estimated to have enough body energy stores in the form of fat and glycogen to last no more than about 1 d under very cold conditions (Alexander et al., 1975; Okamoto et al., 1986; Rowan, 19921. Energy standards are based on the premise that the animal is in a thermoneutral environment during measure- ments of energy transformations. In such an environment, the animal is not required to elicit specific heat-conserving or heat-dissipating mechanisms to maintain core body tem- perature (National Research Council, 19811. The thermo- neutral zone shifts depending on many factors, the more important factors being age, amount of feed intake, amount of subcutaneous fat, and length and thickness of hair coat. The thermoneutral zone in very young calves ranges from 15-25C. Thus, when the environmental temperature drops below 15C, which is referred to as the lower critical temperature, the calf must expend energy to maintain its body temperature. In practical terms, the maintenance energy requirement is increased. For older calves and

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Nutrient Requirements of the Young Calf 221 calves at greater feed intakes, the lower critical tempera- ture may be as low as5 to10C (Webster et al., 19781. Data in Table 10-5 illustrate the effects of a decrease in environmental temperature below the lower critical tem- perature of the calf on energy requirements for mainte- nance. The values were calculated from research data of Schrama (19931. Note in the example given in Table 10-5 that if the lower critical temperature is 10C and the effec- tive ambient temperature is 0C, the maintenance energy requirement is increased by 27 percent. This calculation agrees with experimental findings (Scibilia et al., 19871. Effects of cold stress in increasing maintenance require- ments have been incorporated into the computational model provided with this publication. It is clear from these and other data that calves, especially very young calves, should be fed extra energy during cold weather to satisfy the increase in maintenance energy requirements. That can be accomplished by increasing the amount of liquid diet being fed, by adding additional milk solids to the liquid diet, or by incorporating additional fat into the liquid diet (Schingoethe et al., 1986; Scibilia et al., 1987; [aster et al., 19901. However, additional fat in milk replacer or starter decreases starter intake (Kuehn et al., 1994), which negates at least a portion of the increased energy density from fat supplementation. If additional sol- ids are fed, the DM concentration of milk replacer should not exceed 20 percent to avoid problems with excessive mineral intake (Jenny et al., 1978; Ternouth et al., 1985), and supplemental water should be provided. The availabil- ity of free water is critically important to starter intake (Kertz et al., 19841; provision of warm water 2-3 times daily during cold weather may help to stimulate starter feed intake, which also would help to counteract cold stress. PROTEIN REQUIREMENTS OF CALVES In contrast with the 1978 edition of Nutrient Require- ments of Dairy Cattle (National Research Council, 1978), the 1989 edition provided little information on the protein requirements of young calves weighing less than 100 kg. The tabular data given for protein requirements in the 1989 edition could not be reproduced with information provided (see chapter 9 of Davis and Drackley, 19981. The present edition computes the protein requirement of calves weighing up to 100 kg with the factorial method of Blaxter and Mitchell (19481. The requirement is partitioned into components of maintenance and gain. Maintenance constitutes obligatory nitrogen (N) losses in urine and feces, whereas gain per- tains to N stored in tissues. The protein requirement is expressed in terms of apparent digestible protein (ADP, g/d) and is computed as follows: ADP, g/d = 6.25 L1/BV (E + G + M x D) M x D] (10-4) where BV = biological value (discussed below). Endoge- nous urinary N (E, g/d) is computed as 0.2LW75 (Agricul- tural Research Council, 1980), where live weight (LOO) is in kilograms. This value is somewhat higher than that (0.165 LW075) computed with the formula (2.75 g of net protein per kilogram LW~) given in the 1989 National Research Council publication; however, both are within the range of values in the scientific literature (Blaxter and Wood, 1951; Cunningham and Brisson, 1957; Roy, 19701. The amount of N in gain (G) is assumed to be constant at 30 g N/kg LWG, which is in the range of values reported by others (Blaxter and Wood, 1951; Roy, 1970; Donnelly and Hutton, 1976b; National Research Council, 1978; Davis TABLE 10-5 Effect of Environment on Energy Requirement of Young Calvesa Environmental Increase in Maintenance Energy Maintenance Energy Requirement Increase in ME Temperature Requirement (kcal of NEM/day) (kcal of ME/day)b Required for Maintenance Birth to 3 wk Birth to 3 wk Birth to 3 wk F C of agec >3 wk of aged of agec >3 wk of aged of agec >3 wk of aged 68 20 0 0 1,735 1,735 0 0 59 15 187 0 1,969 1,735 13 0 50 10 373 0 2,203 1,735 27 0 41 5 560 187 2,437 1,969 40 13 32 0 746 373 2,671 2,205 54 27 23 - 5 933 568 2,905 2,437 68 40 14 - 10 1,119 746 3,139 2,671 86 54 5 - 15 1,306 933 3,373 2,905 94 68 - 4 - 20 1,492 1,119 3,607 3,139 108 81 13 - 25 1,679 1,306 3,834 3,373 121 94 22 - 30 1,865 1,492 4,O66 3,607 134 107 Calculated for calf weighing 45.35 kg (100 lbs; 17.35 kg075). Extra heat production = 2.15 kcal/kg075 per day for each degree decrease in environmental temperature below lower critical temperature (Schema, 1993). Because heat production is in terms of net energy (NE), metabolizable energy (ME) was computed as ME = NE/0.8. Maintenance energy requirement 100 kcal/kg075 per day. Calves from birth to 3 wk of age have lower critical temperature in range of 15-25 C. Data above were calculated on basis of lower critical temperature 20 C. Data for calves older than 3 wk of age were calculated on basis of lower critical temperature 10 C.

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222 Nutrient Requirements of Dairy Cattle and Drackley, 19981. Insufficient data were available to describe changes in N content of LW gain as a function of increasing growth rate. Metabolic fecal N (M) is set as 1.9 g/kg of dry matter consumed (D) from milk or milk replacer and 3.3 g/kg of starter DM consumed (Roy, 19801; these values are additive for calves fed both milk and starter. Loss of N in scurf (hair and skin) is ignored in the present edition. The 1989 edition calculated the loss as 0.032 g of N/kg of LW06, which equates to a daily loss of 0.33 g of N for a 50-kg calf. In practice, this loss is compensated by the higher endogenous N losses predicted in the present edition (3.76 g of N for a 50-kg calf than in the 1989 edition (3.10 g of N). The biological value (BV) of milk proteins, equated to the efficiency of N use for growth above maintenance, is assigned a value of 0.80 (Donnelly and Hutton, 1976a). The same factor is assumed to apply for efficiency of use of dietary protein for maintenance functions. This value was determined at limiting protein intakes and assumes that the diet being fed is properly balanced for all essential nutrients and that energy intake is sufficient to support protein synthesis. Protein intake must not be in excess of that required for the targeted gain allowed by energy intake. The BV decreased as protein intake was increased in the studies of Donnelly and Hutton (1976a). The 1978 National Research Council publication used a value of 0.77. Recent studies by Terosky et al. (1997) found that apparent BV for milk replacers containing 21 percent CP from skim milk protein, whey protein concentrate, or mixtures of the two ranged from 0.692 to 0.765. Estimates oftrue biological value (corrected for endogenous N loss and metabolic fecal N) from that study are in excess of 0.80. The conversion of CP to ADP was assumed to be 93 percent for milk proteins (Agricultural Research Council, 1980), which is slightly higher than the value for conversion of dietary CP to absorbable amino acids (91 percent) used in an earlier edition of this publication (National Research Council, 19781. Users should note that requirements for ADP and crude protein (CP) have been established on the basis of diets containing milk proteins with high digestibility and BV; calves might not use alternative, nonmilk proteins in milk replacers at these high efficiencies, and appropriate adjustments may need to be made when such protein sources are used to ensure adequate supply of amino acids for growth (Davis and Drackley, 19981. Furthermore, because digestion of even high-quality milk proteins is immature during the first 2-3 weeks of age (Arieli et al., 1995; Terosky et al., 1997), the value of milk proteins may be overestimated during the early liquid-feeding period. Similar to the situation for energy requirements, however, the subcommittee concluded that information was insuff~- cient to model increasing CP digestibility in the young calf. The BV of absorbed proteins supplied by starter is set at 0.70 (National Research Council, 19781. Calves fed milk plus starter and weaned calves (fed starter only) derive a portion of their protein needs from microbial protein produced in the rumen. However, insufficient data were available to allow calculations of the amounts of rumen- degradable protein (RDP) or rumen-undegradable protein (RUP) supplied with any degree of confidence; thus, the factorial approach using ADP was adopted for calves weighing up to 100 kg. Requirements also are presented in terms of CP. The conversion of CP to ADP is assumed to be 75 percent for starter and grower feeds (Agricultural Research Council, 19801. Quigley et al. (1985) found that an average of 58 percent of the protein reaching the aboma- sum of weaned calves was of microbial origin; flows of N to the abomasum were not reported. Assuming that N flow to the abomasum approximated N intake, that microbial CP is 80 percent true protein that is 80 percent digestible (National Research Council, 1989), and that undegraded feed proteins are 80 percent digestible (National Research Council, 1989) leads to a conversion of CP to ADP of about 71 percent; adoption of the slightly higher value of 75 percent from Agricultural Research Council (1980) leads to better agreement with literature data. The BV and conver- signs of ADP to CP for calves fed starter and milk or starter and milk replacer are assumed to be additive on the basis of the relative amounts of CP supplied by starter and milk (or milk replacer). Examples of requirements for ADP and CP for calves fed milk or milk replacer only, milk replacer plus starter, veal calves, and weaned (ruminant) calves are found in Tables 10-1, 10-2, 10-3, and 10-4, respectively. MINERAL AND VITAMIN REQUIREMENTS OF CALVES Detailed information on the specific roles of mineral elements and vitamins in the nutrition and metabolism of dairy cattle is presented in Chapters 6 and 7. Since the last edition (National Research Council, 1989), there have been few definitive studies of and few problems associated with the field application of the previous recommendations that warrant making major changes in recommendations for most mineral elements or vitamins in Flints of amen 1 _~1 .1 . 1 1 ~ J _ ~ _ _ ~ calves. changes that have been made in the recommenda- tions are discussed below. Minerals The recommended dietary concentrations of mineral elements and vitamins are shown in Table 10-6. For cal- cium and phosphorus recommended concentrations in milk-replacer diets were increased compared with those

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Nutrient Requirements of the Young Calf 223 TABLE 10-6 Mineral and Vitamin Concentrations Recommended for Diets of Young Calves, Compared with Average for Fresh Whole Milk (DM basis) Nutrients Milk Replacerb Starter Feed Grower Feed Whole Milk Minerals Ca (%) 1.00 0.70 0.60 0.95 P (%) 0.70 0.45 0.40 0.76 Mg (%) 0.07 0.10 0.10 0.10 Na (%) 0.40 0.15 0.14 0.38 K (%) 0.65 0.65 0.65 1.12 cl (%) 0.25 0.20 0.20 0.92 S (%) 0.29 0.20 0.20 0.32 Fe (mg/kg) 100C 50 50 3.0 Mn (mg/kg) 40 40 40 0.2-0.4 Zn (mg/kg) 40 40 40 15-38 Cu (mg/kg) 10 10 10 0.1-1.1 I (mg/kg) 0.50 0.25 0.25 0.1-0.2 Co (mg/kg) 0.11 0.10 0.10 0.004-0.008 Se (mg/l~g) 0.30 0.30 0.30 0.02-0.15 Vitamins A (IU/kg of DM) 9,000 4,000 4,000 11,500 D (IU/kg of DM) 600 600 600 307 E (IU/kg of DM) 50 25 25 8 aB-complex vitamins are necessary only in milk-replacer diets. Required concentrations (mg/kg of DM): thiamin, 6.5; riboflavin, 6.5; pyndox~ne, 6.5; pantothenic acid, 13.0; niacin, 10.0; biotin, 0.1; folio acid, 0.5; Bra, 0.07; choline, 1,000. b Required concentrations specified for milk replacer fed at 0.53 kg of DM per day to 45-kg calf. Assuming ME content of 4.75 Mcal/kg, this amount of milk replacer would provide energy-allowable growth of 0.3 kg/d. Concentrations of minerals and vitamins specified will provide adequate daily amounts of minerals and vitamins as defined in Chapters 6 and 7 and in text of this chapter. User is cautioned that feeding larger or smaller amounts of milk replacer, or same amount of milk replacer to larger or smaller calf, changes expected growth arid, consequently, requirements for many vitamins and minerals. CFor veal calves, decrease to less than 50 mg/kg of DM. Of National Research Council (1989), from 0.7 to 1.0 per- cent for calcium and from 0.6 to 0.7 percent for phospho- rus. Recommended concentrations are closer to those found in whole milk (see Table 10-61. Previous calcium recommendations were made considering a fat content in milk replacer of 10 percent, whereas a majority of commer- cial milk replacers today contain fat at 18-22 percent. Higher dietary fat results in increased loss of calcium in the feces because of soap formation between calcium and long-chain fatty acids in the gut (Toullec et al., 19801. The recommended content of sodium in milk replacer was increased from 0.10 to 0.40 percent and from 0.20 to 0.25 percent for chloride (Table 10-61. The committee is unaware of any problems in young calves posed by the previous recommendations for sodium and chloride, but whole milk and most milk replacers that contain milk prod- ucts usually are substantially higher in sodium and chloride than even the new recommendations, thus making practical deficiencies unlikely. As stated earlier, the solids content of milk replacers should be maintained less than 20 percent and free drinking water should be available to avoid prob- lems with excessive intakes of sodium and chloride. The potassium requirement was left unchanged at 0.65 percent of DM for milk-replacer, starter, and grower diets. Well et al. (1988) compared dietary potassium concentra- tions of 0.55, 0.84, 1.02, and 1.32 percent of DM for calves from 4 to 14 wk of age. They detected no differences in feed intake, average daily gain, or mineral status among treatments. In a second trial, Well et al. (1988) compared dietary potassium concentrations of 0.34 and 0.58 percent for calves from 6 to 14 wk of age. Feed intake and live weight gain were greater for calves fed 0.58 percent potass- ium. The authors concluded that potassium requirement of growing dairy calves was "within the range of 0.34 to 0.58 percent," but no concentrations between 0.58 and 0.84 percent were tested. Consequently, the subcommittee concluded that there was insufficient evidence to decrease the requirement from the current value of 0.65 percent of DM. Requirements for most of the trace minerals are unchanged from the previous edition of Nutrient Require- ments of Dairy Cattle (National Research Council, 19891. The required concentrations of iodine were increased from 0.25 mg/kg to 0.50 mg/kg on the basis of information described in Chapter 6, although, as in the situation with sodium and chloride, no indication of deficiency has been noted under practical conditions. The content of cobalt was increased slightly, from 0.10 to 0.11 mg/kg, to be consistent with requirements for other classes of cattle (Chapter 61. Recommended contents of most macromin- Oral elements in milk replacer and starters are close to those of whole milk, whereas recommendations for many of the trace mineral elements are higher than those found in milk, to prevent deficiencies. Caution should be exer- cised in making drastic changes in dietary concentrations of a specific mineral element without being aware of the

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224 Nutrient Requirements of Dairy CattIe possible effects of such changes on the status of other mineral elements (McDowell, 1992~. Vitamins VITAMIN A The subcommittee has markedly increased require- ments for vitamin A in all classes of dairy cattle for reasons discussed in Chapter 7. The requirement for vitamin A in calves was increased from 42.4 (National Research Council, 1989) to 110 IU/kg of LW in the present edition. Eaton et al. (1972) suggested, on the basis of changes in cerebro- spinal fluid pressure, that the requirement for vitamin A should be 96.7 IU/kg of LW for growing Holstein calves. In the Nutrient Requirements of Dairy Cattle (National Research Council, 1989), these data were discussed, but the requirement was not increased; the subcommittee stated that "if substantial evidence for a higher vitamin A requirement is forthcoming, the requirement should be raised." Data from Swanson et al. (2000) demonstrated that an intake of about 134 IU/kg of LW (9,000 IU/kg of DM) maintained liver vitamin A stores in male Holstein calves fed milk replacer, whereas 93 IU/kg or less resulted in decreases in liver concentrations of vitamin A. Calves in that study had received adequate colostrum after birth, were healthy, and were housed under nonstressful environ- mental conditions throughout the study. No clinical mea- sures were affected in that study, even at vitamin A intake (34 IU/kg of LW) less than the previous requirement. However, the liver concentration of vitamin A is believed to be a much more sensitive indicator of vitamin A status than measures used previously to establish requirements. The new requirement was set to be the same as for other classes of cattle and is between the estimates made by Eaton et al. (1972) and Swanson et al. (2000~. Required concentrations have been increased to 9,000 IU/kg of DM for milk replacer and 4,000 IU/kg of DM for starter and grower diets in the present edition. The concentration recommended here for starter or grower feeds will provide required amounts of vitamin A for weaned calves weighing less than 100 kg and gaining 400-900 g/d (Table 10-4~. The presumed safe limit for vitamin A is 66,OOO IU/kg of dietary DM for lactating and nonlactating cattle (National Research Council, 1987), but safe limits specifically for young calves have not been established. Supplementation levels of several times the requirement established in the present edition are common in commercial milk replacers (Tomkins and [aster, 19911. Data to firmly support such a practice are not available. Eicher et al. (1994) found improved fecal consistency in calves fed milk replacer that contained vitamin A at 87,OOO IU/kg, with no effect on vitamin E status. In contrast, several studies have reported adverse effects of high vitamin A on vitamin E status and on other measures of calf health and growth (see Nonnecke et al., 19991. Calves fed a milk replacer containing vitamin A at 44,000 IU/kg of DM rapidly accumulated vitamin A in liver but showed no signs of toxicity during 28 days of feeding (Swanson et al., 2000~. Supplementation with vitamin A in amounts greater than recommended in the present edition cannot be justified on the basis of available data. In particular, caution should be observed in formula- tion of milk replacers for veal calves and for replacement calves in accelerated-growth schemes to avoid potential problems with excessive vitamin A intake. VITAMIN E The requirement for vitamin E for calves continues to be debated. Requirements for vitamin E were increased substantially for lactating and dry cows in the present edi- tion (Chapter 7~. The subcommittee has increased the requirement for vitamin E for calves by 25 percent, from 40 IU/kg of dietary DM to 50 IU/kg. The decision to increase the vitamin E requirement represents a compro- mise until more-def~nitive data are available. The increase is based on two main factors. First, although 40 IU/kg of DM is adequate to prevent classic signs of deficiency, such as muscular dystrophy or retardation of growth of calves in controlled systems, calves under conditions of stress more typical in practice might require higher intakes of vitamin E to augment the immune system. Vitamin E sup- plementation improved immune-system responses, as mea- sured by lymphocyte stimulation indexes, IgM concentra- tions, serum cortisol concentrations, and antibody response to a booster vaccine (Ready et al., 1986, 1987b). Indicators of cell-membrane damage (serum creatine kinase, glutamic oxalacetic transaminase, and lactic acid dehydrogenase) suggested that Vitamin E supplementation protected mem- branes from oxidative damage (Ready et al., 1986, 1987b). Vitamin E functions as an antioxidant and interacts with selenium to maintain the structural integrity of tissues (Combs, 1992; McDowell, 19921. Reddy et al. (1987a) suggested on the basis of a study in which calves were supplemented with 125, 25O, or 500 IU of vitamin E per day that the requirement was about 2.4 IU/kg of body weight. However, no supplementation levels lower than 125 IU/d were tested, and numbers of animals were insufficient to determine clinical responses. The subcommittee felt that, in the absence of large-scale dose-response studies to determine clinical responses, such a large increase was not justified. Furthermore, increased requirements for dry cows should increase concentrations of vitamin E in colostrum (Quigley and Drewry, 1998), which could provide more vitamin E to calves than was consumed by calves in the Kansas State University studies. Second, the relationship of vitamin E with other dietary nutrients must be considered. For the young calf, dietary

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Nutrient Requirements of the Young Calf 225 vitamin E should be balanced with the content of essential fatty acids (1.5-2.5 IU of vitamin E per gram of linoleic acid; Stobo, 1983) to prevent oxidative stress from increased intake of polyunsaturated fatty acids, as in young nonruminant animals. With typical daily intakes of 10-15 g of linoleic acid from milk replacers, 15-38 IU of vitamin E daily would be necessary, according to guidelines of Stobo (19831. To supply adequate vitamin E to meet this guideline for a calf fed 600 g of milk-replacer DM daily, vitamin E content would need to be 25-63 IU/kg of DM. Some evidence suggests that increased vitamin A in the diet decreases the bioavailability of vitamin E (see Non- necke et al., 19991. Consequently, the moderate increase in the vitamin E requirement also is justified because of the substantially increased vitamin A requirement. Diar- rhea and gut infections decrease fat digestion and hence lower the absorption of the fat-soluble vitamins A, D, and E. Given the widespread occurrence of digestive distur- bances in young calves before weaning, the increases in recommendations for both vitamin A and vitamin E should be beneficial in practical situations. The subcommittee recognizes that the requirement for vitamin E might need to be adjusted in future editions if data from large-scale dose-response studies become available. VITAMIN D AND WATER-SOLUBLE VITAMINS Requirements for vitamin D were not changed from the 1989 edition (Table 10-61. Water-soluble vitamins must be included in the milk-replacer diet of calves (see Table 10- 61. Once the calf is weaned to dry feed, there is no evidence that these vitamins need to be supplemented to the diet, inasmuch as the microorganisms in the digestive tract syn- thesize ample amounts to meet the needs of the calf. FEED-COMPOSITION DATA WITH APPLICATION TO DIET FORMULATIONS FOR CALVES Values for digestible energy and metabolizable energy for foodstuffs for calves in the National Research Council (1989) are realistic compared with known gross energy and digestibility data and agree closely with values assigned by other sources. However, as pointed out earlier in this chapter, the NEM and NEG values for milk, milk byprod- ucts, and milk replacers given in the 1989 edition were too low according to reported efficiencies of use of ME by young milk-fed calves (see chapter 9 in Davis and Drackley, 19981. The problem arose from the inappropriate use of the equations derived by Garrett (1980) from growth stud- ies with feedlot cattle to derive the net energy of liquid diets for nonruminant calves. A different approach has been taken in the present edition to establish the net energy values for calf diets. Gross energy (GE) values have been calculated from data on composition and heat of combustion. For milk and milk-derived ingredients used in milk replacers, GE (Meal/kg) = 0.057 CP% + 0.092 fat% + 0.0395 lactose%, (10-5) where lactose was calculated as 100CP%fat%ash%; all components are expressed on a DM basis. For whole milk, milk replacers, and milk-derived ingredients, DE was calculated as 0.97 GE. For all milk and milk products, including milk replacers, ME was calculated as 0.96 DE. Values calculated by these methods agree closely with those in the previous edition of this publication (National Research Council, 19891. The NEM values for milk, milk-derived ingredients, and milk replacers is calculated as 0.86 ME, consistent with the NEM requirements discussed earlier. This is similar to the value of 0.85 used by the Agricultural Research Council (19801. The approach used to derive values for NEG for milk and milk-derived ingredients is based on the relation- ship between the metabolizability (q) of the diet (ME/GE) and the efficiency of use of ME for maintenance and gain (Agricultural Research Council, 19801. The NEGvalues for milk-based diets can then be estimated as follows (Agricul- tural Research Council, 19801: NEG = (0.38q + 0.337) ME (10-6) Values for q have been computed and are included in Table 10-7, which provides composition data for ingredi- ents used in milk replacers. The values for NEM and NEG calculated by these methods agree well with efficiencies of use of ME of 80 and 69 percent for maintenance and gain, respectively, determined by others (Roy, 1980; Toul- lec, 19891. A slightly different procedure was used to calculate NEM and NEG values for ingredients used in starter and grower diets. For all nonmilk ingredients, GE (Meal/kg) = 0.057 CP% + 0.094 ether extract (EE)% + 0.0415 carbohydrate% (10-7) where carbohydrate was calculated as 100CP%fat% ash%. The DE values were calculated as the sum of the products of digestible CP, EE, and carbohydrates multiplied by their heats of combustion; this is the approach described in Chapter 2 to calculate energy values for feeds fed to other classes of dairy cattle in this edition. Values for ME were calculated with the approach in the previous edition (National Research Council, 1989), except that the equation was corrected to reflect increased eff~- ciency of use of fat:

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226 Nutrient Requirements of Dairy Cattle TABLE 10-7 Energy, Protein, Calcium, and Phosphorus Concentrations in Feedstuffs Commonly Used in Formulation of Milk Replacers for Young Calvesa GE DE ME NEM NEG CP EE Ca P Ash International DM (Meal/kg (Mcal/l~g (Mcalikg ME/GE (Meal/kg (Mcal/l~g Feed Feed Number (%) of DM) of DM) of DM) (q) of DM) of DM) % of DM Whole milk 5-01-168 12.5 5.76 5.59 5.37 0.93 4.62 3.70 25.4 30.8 1.00 0.75 6.3 Skim milk, fresh 5-01-170 10 4.31 4.19 4.02 0.93 3.46 2.77 35.5 0.3 1.35 1.02 6.9 Skim milk, powder 5-01-175 94 4.38 4.25 4.08 0.93 3.51 2.82 37.4 1.0 1.29 1.08 6.9 Whey-powder 4-01-182 93 3.92 3.80 3.65 0.93 3.14 2.52 13.5 1.0 0.76 0.68 8.1 Whey protein concentrate 93 4.48 4.35 4.17 0.93 3.59 2.88 37.1 2.2 0.54 0.60 6.0 Whey, fresh 4-08-134 7 3.89 3.78 3.62 0.93 3.12 2.50 14.2 0.7 0.73 0.65 8.7 Whey, delactosed 4-01-186 93 3.65 3.54 3.40 0.93 2.92 2.34 17.9 0.7 1.71 1.12 16.5 Whey permeate 98 3.66 3.55 3.41 0.93 2.93 2.35 3.7 0 1.77 0.97 9.0 Casein 5-01-162 91 5.45 5.29 5.08 0.93 4.37 3.50 92.7 0.7 0.40 0.35 4.0 aData from NRC (1989); Toullec (1989); Tomkins and Jaster (1991). Calculations are described in text. ME = (1.01 x DE 0.45) + 0.0046 (EE 3) (10-8) where ME and DE are Mcal/kg and EE is percent of dietary DM. These ME values are analogous to ME values at maintenance for older cattle (Chapter 2) and are more consistent with known efficiencies of conversion of DE to ME, given that methane production in young calves is extremely low (Gonzlalez-;[imenez and Blaxter, 1962; Holmes and Davey, 1976). Values for NEM and NEGwere calculated as described in the section on energy require- ments earlier in this chapter. For NEM and NEG, ME as calculated above was multiplied by the respective eff~cienc- ies of 0.75 for maintenance and 0.57 for gain. These eff~- ciencies are similar to those estimated by others from the metabolizability (q) of ingredients. For example, Agricul- tural Research Council (1980) calculated NEM as (0.287q + 0.554)ME and NEG as (0.78q + 0.006)ME. For a calf starter with q = 0.7O, efficiencies for maintenance and gain would be 75 and 55 percent when calculated with the Agricultural Research Council equations. Table 10-8 presents composition data on examples of three typical milk replacers, a starter diet, and a grower diet for calves. The values presented for NEM and NEG content are considerably higher for all feeds than those calculated with the previous methods (National Research Council, 1989). The computer model automatically calcu- lates ME, NEM, and NEGconcentrations for feeds used for young calves. Users are cautioned that the requirements and feed values are designed to be used together. Use of NEM and NEG values from previous editions with the pres- ent growth model, or vice versa, will result in erroneous predictions. Values for total digestible nutrients (TDN) are not given for calf requirements or feeds in this edition. If desired, TDN can be calculated as described for feeds for other classes of cattle (see Chapter 2). For milk, milk replacer, and milk ingredients, TDN = 0.93 CP + (EE x 2.25) + 0.98 (100CPEEAsh) - 7 (10-9) OTHER ASPECTS OF CALF NUTRITION Fetal Nutrition Although severe undernutrition can impair normal fetal development (National Research Council, 1968), the developing fetal calf is afforded a high priority for maternal nutrients. Moderate underfeeding of either protein or energy did not result in measurable changes in calf birth weight, viability, or health (Davis and Drackley, 1998; Quigley and Drewry, 1998). Prolonged restriction of pro- tein or energy during gestation decreased thermogenic abilities of beefcalves at birth (Carstens et al., 1987; Ridder et al., 1991). Maternal deficiencies of phosphorus, manganese, cobalt, copper, zinc, and selenium can result in deficiencies in the fetus and newborn calf (National Research Council, 1968). The fetus has the ability to concentrate some of these minerals, particularly copper (Hidiroglou and Knipfel, 1981) and selenium (Van Saun et al., 1989a), providing some protection against marginal deficiencies in the mother. Selenium supplementation of pregnant cows increased selenium reserves in the newborn calves (Abdel- rahman and Kincaid, 1995). Placental transfer of vitamin E to the developing fetus is low, although the fetal calf appears to have some ability to concentrate vitamin E from the dam (Van Saun et al., 1989b). The calf is born with a low vitamin E status and is highly dependent on intake of colostrum and then milk or milk replacer to obtain needed vitamin E during early postnatal life. If diets for pregnant cows are balanced to meet recommendations for pregnancy and maternal growth (see Chapters 6 and 7), as well as for optimal transition success (see Chapter 9), nutrient supply should be adequate for normal growth and development

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Nutrient Requirements of the Young Calf 227 TABLE 10-8 Energy, Protein, Fiber, and Mineral Composition of Three Milk Replacers (MR), a Starter Feed, and a Grower Feed for Young Calves GEa DEa MEa NEM NEG (Meal/kg (Meal/kg (Meal/kg (Meal/kg (Meal/kg CP EE ADF NDF Ca P Feed DM) of DM) of DM) of DM) of DM) (%) (%) (%) (%) (%) (%) MR-1 4.61 4.47 4.29 3.69b 2.96C 22 10 1.0 0.70 MR-2 5.10 4.95 4.75 4.09b 3.28C 20 20 1.0 0.70 MR-3 5.07 4.91 4.72 4.o6b 3.26C 18 20 1.0 0.70 Starter 4.49 3.69 3.28 2.46d l.78e 18 3 11.6 12.8 0.7 0 45 Grower 4.36 3.65 3.24 2.43d 1.6le 16 3 8.0 18.0 0.6 0.40 a energy values calculated as follows: Gross energy (GE) is calculated from composition and heat of combustion. For milk replacers, GE (kcal/kg) = grower, GE (kcal/kg) = 0.057 CP + 0.092 EE + 0.0415 carbohydrate. For MR, digestible energy (DE) = 0.97 GE. For starter and grower, DE is calculated as sum of digestible protein, fat, and carbohydrates, each multiplied by heat of combustion. For MR, metabolizable energy (ME) calculated as 0.93 GE (ME/GE of whole milk has been measured at 0.93; Roy, 1980). For starter and grower feeds, ME = (1.01 X DE0.45) + (0.0046EE 3) (see text and Chapter 2). bNEM = 0.86 ME. See text for details. CNEG = (0.38q + 0.337) X ME. Based on q of 0.93 y~elding an eff~ciency of 0.69 for ME use (ARC, 1980). ~NEM = ME X 0.75. eNEG = ME X 0.57. - 0.057 CP + 0.092 fat + 0.0395 lactose. For starter and of the fetal calf (Davis and Drackley, 1998; Quigley and Drewry, 1998). Colostrum Calves are born with negligible circulating concentra- tions of immunoglobulins (McCoy et al.,19701. Early provi- sion of high-quality colostrum in amounts suff~cient to pro- vide at least 100 g of IgG is critical to calf survival and well-being (Davis and Drackley, 1998; Quigley and Drewry, 19981. The immunoglobulin content of colostrum is highly variable (Pritchett et al., 19911; therefore, to max- imize the likelihood of obtaining suff~cient IgG, it is recom- mended that calves be fed at least 3 L of colostrum from multiparous cows within an hour afterbirth. Holstein calves can be administered as much as 3.8 L of colostrum in a single feeding after birth to ensure delivery of suff~cient IgG (Besser et al., 1991; Hopkins and Quigley, 19971. In addition to disease protection, earlyprovision of colos- trum is important as a source of nutrients (Davis and Drackley, 1998; Quigley and Drewry, 19981. Because sup- plies of endogenous fuels are exhausted within hours with- out feed (Okamato et al., 1986; Rowan, 1992), the carbohy- drate, fat, and protein in colostrum are essential as fuels for the newborn. Most of the essential minerals and vita- mins are substantially more concentrated in colostrum than in milk (Foley and Otterby, 19781. Consumption of ade- Water and Electrolytes quate amounts of colostrum by the newborn calf, followed by consumption of milk or milk replacer that is adequate in mineral and vitamin content, is important to compensate for any maternal inadequacies during gestation. Increasing evidence in calves and other species indicates that colos- trum also provides a number of hormones and growth factors necessary to stimulate growth and development of the digestive tract and other organ systems (Hammon and Blum, 19981. Commercial products containing immunoglobulins may be useful to supplement poor-quality colostrum (Gerry et al., 1996; Morin et al., 1997; Arthington et al.,20001. Other products are designed to be injected to increase serum immunoglobulins in calves (Quigley and Welborn, 19961. At present, none of the commercially available supple- ments or substitutes can completely replace colostrum in providing passive immunity to calves (Arthington et al., 20001. High-quality colostrum should be provided when- ever possible; supplements are of little additional value when sufficient amounts of high-quality colostrum are administered (Hopkins and Quigley, 19971. Development of products that can deliver suff~cient biologically active immunoglobulins to the newborn calf might be increasingly important for use in biosecurity programs to control conta- gious diseases, such as ;[ohne's disease, in which it would be desirable to avoid the feeding of any colostrum or whole milk to calves. Although the nutritional aspects of colos- trum probably could be replaced by a properly formulated milk replacer, the consequences of the absence of the growth factors and hormones normally consumed in colos- trum are not known. Water is the most important nutrient and, although essential, is often overlooked. Too often, it is assumed that if a calf is being fed a liquid diet, its needs for water will be satisf~ed. Fresh water, in addition to water consumed as part of the diet, is essential for optimal growth and consumption of dry feed (Leaver and Yarrow, 1972; Kertz et al., 19841.

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228 Nutrient Requirements of Dairy CattIe Aside from constituting 70-75 percent of the weight of the calf, water plays important roles as a solvent for nutri- ents, a thermoregulator, and an osmoregulator (Davis and Drackley, 19981. Calves, because of their greater propen- sity to develop digestive disturbances (diarrhea), experi- ence greater problems with water balance than do older animals. During incidents of diarrhea, 10-12 percent of body weight can be lost as water. The water loss in feces carries with it major losses of the electrolytes sodium, chloride, and potassium (Lewis and Phillips, 1978; Phillips, 19851. Such losses of water and electrolytes result in severe dehy- dration and electrolyte imbalances, which if not rapidly corrected will result in death. In fact most deaths associated with diarrhea occur from these phenomena rather than directly from infectious agents (Booth and Naylor, 19871. Recent evidence indicates that electrolyte disturbances are more important than dehydration itself in causing death from diarrhea (Walker et al., 19981. At the first signs of diarrhea, a calf should be started on oral rehydration (Davis and Drackley, 19981. Current information suggests that the calf should continue to receive a portion of, if not all, its regular feeding of milk or milk replacer with the oral electrolyte product (McGuirk, 1992; Garthwaite et al., 1994) as long as it is alert and willing to drink. Calves that are severely dehydrated, recumbent, or acidemic will require intravenous fluid ther- apy for recovery. Milk Replacers Milk replacers are used on a majority of dairy farms in the United States (Heinrichs et al., 19951. Substantial changes in milk-replacer formulation have occurred since the last edition of this publication (National Research Council, 19891. Increases in market prices for dried skim milk, coupled with development of low-temperature ultra- f~ltration techniques for preparation of high-quality whey protein concentrates, have led to the almost complete replacement of dried skim milk with whey-derived prod- ucts (Davis and Drackley, 19981. Milk-replacer formula- tions generally are classified as all-milk protein or as alter- native protein. Milk replacers of all-milk protein contain whey protein concentrate, dried whey, and delactosed whey as protein sources. Many alternative-protein formula- tions are available, in which portions of the milk proteins (typically 50 percent) are replaced with lower-cost ingredi- ents, such as soy protein concentrate, soy protein isolates, animal plasma or whole-blood proteins, and modified wheat gluten (Davis and Drackley, 19981. Examples of formulations and a review of recent research can be found in chapter 14 of Davis and Drackley (19981. Aspects of milk replacer use also have been reviewed by Heinrichs (1994, 19951. The ability of these protein sources to supply an ade- quate amount and profile of amino acids for growth of preruminant calves depends on the amino acid profile of the protein, the quality of the manufacturing process, and the ability of the calf to digest the protein. High tempera- tures during drying can damage proteins and lessen their biologic value (Wilson and Wheelock, 19721. Furthermore, antinutritional factors present in some protein sources can decrease efficiency of amino acid use (Huisman, 1989; Lalles, 19931. Whey protein concentrate is digested and utilized as least as well as skim milk protein by young calves (Terosky et al., 1997; Lammers et al., 19981. The proteolytic digestive system of the young calf is immature at birth, and until the age of about 3 weeks the calf is less able to digest most nonmilk proteins (Toullec and Guilloteau, 19891. Therefore, for optimal growth dur- ing the first 3 weeks of life, it is recommended that milk replacers containing only milk proteins be used. Older calves are able to use formulations that contain nonmilk proteins. Milk replacers typically contain tallow, choice white grease, or lard as a fat source. The degree of homogeniza- tion is critical for high digestibility (Raven, 19701. Emulsifi- ers, such as lecithin and monoglycerides, often are added to enhance mixing characteristics and fat digestibility. In general, vegetable oils and fat sources that contain large amounts of free fatty acids are poorly used by calves (;[en- kins et al., 19851. Research data on optimal concentrations of fat in milk replacers are conflicting, with little definitive evidence that a fat content beyond 10-12 percent is needed, at least in moderate environments (Heinrichs, 19951. Feed Additives A variety of feed additives have been examined for inclu- sion in milk replacers or dry feeds (Heinrichs, 19931. The addition of medications to milk replacers in the US is regulated by the Food and Drug Administration. Antibiot- ics such as oxytetracycline and neomycin are widely used in milk replacers (Heinrichs et al., 19951. Antibiotics consis- tently improve growth rates and feed efficiency and decrease incidence and severity of scouring of calves (Mor- rill et al., 1977; Quigley et al., 1997a), although the mode of action still is poorly understood. Benefits of antibiotic inclusion may be more evident for calves raised intensively in large numbers, for shipped-in calves originating from different farms, and for calves raised under conditions of stress (Morrill et al., 1977; Morrill et al., 1995; Davis and Drackley, 19981. Lasalocid and decoquinate added to feeds are effective in control of coccidiosis (Hoblet et al., 1989; Heinrichs et al., 1990; Heinrichs and Bush, 1991; Eicher-Pruiett et al., 1992; Quigley et al.,1997b). Supplementation in calf starter

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Nutrient Requirements of the Young Calf 229 requires adequate feed intake to achieve effective dosages, but infection with coccidia often occurs before starter intake is sufficient (Quigley et al., 1997b). Bacterial probi- otic products have shown some benefit in improving calf health and performance (Jenny et al., 1991; Higginbotham and Bath, 1993; Morrill et al., 1995; Abe et al., 1995; Cruywagen et al., 1996) although responses have been variable and inconsistent (Morrill et al., 19771. Experimen- tal results from additions of fungal (Beharka et al., 1991) or yeast (Quigley et al., 1992) culture products to starter diets have been inconclusive. Sodium bicarbonate increased starter intake and growth of young calves in one study (Curnick et al., 1983) but did not affect intake or calf performance in another study (Quigley et al., 19921. Practical Feeding Considerations As mentioned in the introduction to this chapter, female calves in the United States destined for herd replacements should be fed restricted amounts of milk or milk replacer (typically 8-10 percent of birth weight) to encourage early consumption of calf starter (National Research Council, 19891. Development of early starter intake is inversely proportional to the amount of liquid fed (Hodgson, 19711. Growth rates of young calves during the liquid feeding period thus are much lower than the maximal growth rates of calves (Khouri and Pickering, 1968; Hodgson, 1971), and feed efficiency is lower than that in the young of other farm animals that consume milk ad libitum (Khouri and Pickering, 1968; Davis and Drackley, 19981. Nevertheless, restricted liquid feeding encourages earlier starter intake and ruminal development, which in turn allows for earlier weaning and more economic body weight gains. Ad libitum or increased liquid feeding programs researched to date have resulted in greater growth rates and improved feed efficiency during the liquid feeding period, but lower con- sumption of dry feed and variable effects on calf health (Khouri and Pickering, 1968; Hodgson, 1971; Huber et al., 1984; Nocek and Braund, 1986; Richard et al., 19881. Methods to capitalize on the early growth potential are being researched in the context of accelerated rearing pro- grams for heifers that encompass all stages of growth from birth to first calving. However, these programs are still under development and evaluation, and cannot yet be rec- ommended at this time. During the early liquid feeding period, growth of calves fed milk or milk replacer is directly proportional to the amount of liquid provided (Khouri and Pickering, 1968; Hodgson, 1971; Huber et al., 19841. In contrast, in restricted liquid feeding programs, growth rates are directly proportional to the amount of calf starter con- sumed (Kertz et al., 1979, 19841. Users should be aware that typical milk replacers contain 10-20 percent less energy than comparable volumes of whole milk because of the lower fat content of milk replacers. A 40-kg calf fed milk replacer at 9 percent of body weight would consume 454 g of DM. If the milk replacer contains ME at 4.7 Mcal/kg of DM, the calf would consume enough energy for maintenance and a body weight gain of 234 g/d under thermoneutral conditions. According to the model pre- sented in this edition, feeding the same volume of whole milk would support a gain of 331 g/d. In contrast, if the same calf is housed at 20C below its lower critical tempera- ture, 454 g/d of milk replacer powder is insufficient even for maintenance. Increasing evidence suggests that these low feeding rates also are inadequate to support optimal health and function ofthe immune system, especially under adverse environmental conditions (Williams et al., 1981; Griebel et al., 1987; Pollock et al., 1993, 19941. High intakes of milk or milk replacer are important for veal production. The effects of increasing intake of whole milk and milk replacer for a 40-kg calf are illustrated in Figure 10-1. Note that the difference in growth perfor- mance predicted between whole milk and milk replacer fed at equal amounts is accounted for entirely by the 13 percent greater ME content of whole-milk solids versus the milk replacer solids. Gains predicted here agree closely with literature studies with high rates of milk feeding (Khouri and Pickering, 1968; Hodgson, 19711. Large-breed calves can be weaned easily when consum- ing at least 0.68 kg of a good-quality starter daily for 3 1200 1 1 _ 1 1 00 - 1 000 - 900 - 800- 700- ._ ~ 400- 600 - 500 - 300 - 200 - 100 - o- ............... ............................... ............................... ............................... ............................... L :::::: :::::::::::::::::::: ............................... ............................... ............................... ............................... . . . ............................... l . . . ............................... l ............................... l ............................... l ............................... l ............................... l ............................... l ............................... l ............................... l ............................... l ............................... l ............................... l ............................... l ............... , .............................. ............................... ............................... ............................... ............................... 10 14 18 Feeding rate (% of BW) FIGURE 10-1 Example of growth rate predicted by the model in this edition for a 40-kg calf fed whole milk (open bars) or milk replacer (dark bars) at 10, 14, or 18 percent of body weight. Whole milk contains ME at 5.37 McaLkg of DM. Milk replacer contains ME at 4.75 McaLkg of DM and is assumed to be reconsti- toted to 12.5 percent solids, similar to total solids content of whole milk.

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230 Nutrient Requirements of Dairy Cattle consecutive days. Under good management, with restricted milk or milk replacer feeding this can occur as early as the age of 4 weeks (Kertz et al., 1979, 19841. More aggressive milk-feeding programs will delay development of starter intake and weaning age (Hodgson, 1971; Huber et al., 19841. Other factors important for early development of d~y-feed intake include free access to supplemental water; provision of palatable starter feeds (generally of coarse texture rather than finely ground); keeping feeds fresh, dry, and free of mold; and good health of calves. A major metabolic factor could be the establishment of stable ~umi- nal fermentation with pH greater than 5.5 (Williams and Frost, 19921. R E F E R E N C E S Abdelgadir, I. E. O., and J. L. Morrill. 1995. Effect of processing sorghum grain on dairy calf performance. J. Dairy Sci. 78:2040-2046. Abdelgadir, I. E. O., J. L. Morrill, and J. J. Higgins. 1996a. Effect of roasted soybeans and corn on performance and ruminal and blood metabolites of dairy calves. J. Dairy Sci. 79:465-474. Abdelgadir, I. E. O., J. L. Morrill, and J. J. Higgins. 1996b. Ruminal availabilities of protein and starch: effects on growth and ruminal and plasma metabolites of dairy calves. J. Dairy Sci. 79:283-290. Abdelrahman, M. M., and R. L. Kincaid. 1995. Effect of selenium supple- mentation of cows on maternal transfer of selenium to fetal and new- born calves. J. Dairy Sci. 78:625-630. Abe, F., N. Ishibashi, and S. Shimamura. 1995. Effect of administration of bif~dobacteria and lactic acid bacteria to newborn calves and piglets. J. Dairy Sci. 78:2838-2846. Agricultural Research Council. 1980. The Nutrient Requirements of Ruminant Livestock. Farnham Royal, Slough, England: Common- wealth Agricultural Bureaux. Akayezu, J. M., J. G. Linn, D. E. Otterby, W. P. Hansen, and D. G. Johnson. 1994. Evaluation of calf starters containing different amounts of crude protein for growth of Holstein calves. J. Dairy Sci. 77:1882-1889. Alexander, G., J.W. Bennett, and R.T. Gammell. 1975. Brown adipose tissue in the newborn calf (Bos Taurus). J. Physiol. 244: 223-234. Anderson, K. L., T. G. Nagaraja, and J. L. Morrill. 1987a. Ruminal metabolic development in calves weaned conventionally or early. J. Dairy Sci. 70:1000-1005. Anderson, K. L., T. G. Nagaraja, J. L. Morrill, T. B. Avery, S. J. Galitzer, and J. E. Boyer. 1987b. Ruminal microbial development in convention- ally or early weaned calves. J. Anim. Sci. 64:1215-1226. Arieli, A., J. W. Schrama, W. Van Der Hel, and M. W. A. Verstegen. 1995. Development of metabolic partitioning of energy in young calves. J. Dairy Sci. 78:1154-1162. Arthington, J. D., M. B. Cattell, and J. D. Quigley, III. 2000. Effect of dietary IgG source (colostrum, serum, or milk-derived supplement) on the efficiency of Ig absorption in newborn Holstein calves. J. Dairy Sci. 83:1463-1467. Beharka, A. A., T. G. Nagaraja, and J. L. Morrill. 1991. Performance and ruminal function development of young calves fed diets with Aspergillus oryzae fermentation extract. J. Dairy Sci. 74:4326-4336. Beharka, A. A., T. G. Nagaraja, J. L. Morrill, G. A. Kennedy, and R. D. Klemm. 1998. Effects of form of the diet on anatomical, microbial, and fermentative development of the rumen of neonatal calves. J. Dairy Sci. 81:1946-1955. Besser, T. E., C. C. Gay, and L. Pritchett. 1991. Comparison of three methods of feeding colostrum to dairy calves. J. Am. Vet. Med. Assoc. 198:419-422. Blaxter, K. L. and H. H. Mitchell. 1948. The factorization of the protein requirements of ruminants and of the protein value of feeds, with particular reference to the significance of the metabolic fecal nitrogen. J. Anim. Sci. 7: 351-372. Blaxter, K. L., and W. A. Wood. 1951. The nutrition of the young Ayrshire calf.4. Some factors affecting the biological value of protein determined by nitrogen balance methods. Br. J. Nutr. 5:55-67. Booth, A. J., and J. M. Naylor. 1987. Correction of metabolic acidosis in diarrhea! calves by oral administration of electrolyte solutions with or without bicarbonate. J. Am. Vet. Med. Assoc. 191: 62-68. Brownlee, A. 1956. The development of rumen papillae in cattle fed on different diets. Br. Vet. J. 112:369-375. Carstens, G. E., D. E. Johnson, M. D. Holland, and K. G. Odde. 1987. Effects of prepartum protein nutrition and birth weight on basal metab- olism in bovine neonates. J. Anim. Sci. 65:745-51. Chester-Jones, H., D. M. Ziegler, and J. C. Meiske. 1991. Feeding whole or rolled corn with pelleted supplement to Holstein steers from weaning to 190 kilograms. J. Dairy Sci. 74:1765-1771. Combs, G. F., Jr. 1992. The Vitamins: Fundamental Aspects in Nutrition and Health. San Deigo, CA: Academic Press, Inc. Cruywagen, C. W., I. Jordaan, and L. Venter.1996. Effect of Lactobacillus acidophilus supplementation of milk replacer on preweaning perfor- mance of calves. J. Dairy Sci. 79:483-486. Cunningham, H. M., and G. J. Brisson. 1957. The endogenous urinary and metabolic fecal nitrogen excretions of newborn dairy calves. Can. J. Anim. Sci. 37:152-156. Curnick, K. E., L. D. Muller, J. A. Rogers, T. J. Snyder, and T. F. Sweeney. 1983. Addition of sodium bicarbonate to calf starter rations varying in protein percent. J. Dairy Sci. 66:2149-2160. Davis C. L., and J. H. Clark. 1981. Ruminant digestion and metabolism. Dev. Ind. Microbiol. 22:247-259. Davis, C. L. and J. K. Drackley. 1998. The Development, Nutrition, and Management of the Young Calf. Iowa State University Press, Ames, Iowa. Donnelly, P. E., and J. B. Hutton. 1976a. Effects of dietary protein and energy on the growth of Friesian bull calves. I. Food intake, growth, and protein requirements. N.Z. J. Agric. Res. 19: 289-297. Donnelly, P. E., and J. B. Hutton. 1976b. Effects of dietary protein and energy on the growth of Friesian bull calves. II. Effects of level of feed intake and dietary protein content on body composition. N.Z. J. Agric. Res. 19: 409-414. Eaton, H. D., J. E. Rousseau, Jr., R. C. Hall, Jr., H. I. Prier, and J. J. Lucas. 1972. Reevaluation of the minimum vitamin A requirement of Holstein male calves based upon elevated cerebrospinal fluid pressure. J. Dairy Sci. 55:232-237. Eicher, S. D., J. L. Morrill, F. Blecha, C. G. Chitko-McKown, N. V. Anderson, and J. J. Higgins. 1994. Leukocyte functions of young dairy calves fed milk replacers supplemented with vitamins A and E. J. Dairy Sci. 77:1399-1407. Eicher-Pruiett, S. D., J. L. Morrill, T. G. Nagaraja, J. J. Higgins, N. V. Anderson, and P. G. Reddy. 1992. Response of young dairy calves with lasalocid delivery varied in feed sources. J. Dairy Sci. 75:857-862. Flatt, W. G., R. G. Warner, and J. K. Loosli. 1958. 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