5
Nutrient Requirements of the Hamster

Taxonomically hamsters are classified as a subfamily, Cricetinae, with 7 genera and 18 species in the family Muridae (Musser and Carleton, 1993). They are distributed throughout the Palearctic zone of Eurasia (Anderson and Jones, 1984). Original habitats of laboratory hamsters included clay deserts, shrub-covered plains, forested steppes, and/or cultivated fields.

Golden hamsters, Mesocricetus auratus, collected from a burrow 8 feet deep in a wheat field near Aleppo, Syria, were established as a colony of laboratory animals at the Microbiological Institute of Jerusalem in 1930. These animals were used to complete research on kala-azar delayed by the failure of Chinese hamsters to breed in captivity (Adler and Theodor, 1931). Adler took breeding pairs to Paris and London to establish colonies at research institutions there (Bruce and Hindle, 1934; Adler, 1948; Murphy, 1985). Breeding stock were distributed to investigators in India, Egypt, and the United States (Doull and Megrall, 1939; Poiley, 1950). Golden hamsters is the hamster species most frequently used in research (Hoffman et al., 1968; Siegel, 1985; Van Hoosier and McPherson, 1987), but very little is known about their nutritional requirements.

In the past 20 years, 7 additional hamster species have been used as laboratory animals. Animals identified as strain MHH:EPH are maintained in Hannover, Germany (Reznik et al., 1978; Mohr and Ernst, 1987). Mouse-like Chinese striped hamsters, Cricetulus barabensis, are used in research in cytogenetics, diabetes, and toxicology (Calland et al., 1986; Diani and Gerritsen, 1987). The large guinea-pig-like European hamster, Cricetus cricetus, formerly considered a pest in agricultural areas, is now a model for research in carcinogenesis. Dwarf hamsters, Phodopus campbelli and P. sungorus, of southern and western Siberia, are used in research in cytogenetics, carcinogenesis, diabetes mellitus, obesity, photoperiod changes, and social behavior (Pogosianz and Sokova, 1967; Hoffmann, 1973; Daly, 1975; Gamperl et al., 1978; Hoffmann, 1978. Steinlechner et al., 1983; Wade and Bartness, 1984; Pond et al., 1987; Ruf et al., 1991). Turkish hamsters, Mesocricetus brandti, are used in hibernation, taxonomy, and cytogenetics research (Lyman and O'Brien, 1977; Lyman et al., 1981, 1983; Todd et al., 1972). Colonies of the Romanian hamster, Mesocricetus neutoni , were established in Bucharest and used for research in cytogenetics and taxonomy (Hamar and Schutowa, 1966; Murphy, 1977; Popescu and DiPaolo, 1980). The Armenian, or migratory hamster, Cricetulus migratorius , is used on a limited basis in cytogenetics and oncology research (Lavappa and Yerganian, 1970; Cantrell and Padovan, 1987) (see Table 5-1).

BIOLOGICAL AND BEHAVIORAL CHARACTERISTICS

Unlike simple-stomached rats, mice, and guinea pigs, hamsters, like voles, have a stomach that consists of two distinct compartments: a keratinized, nonglandular forestomach (cardiac) separated from a glandular region (pyloric) by sphincter-like muscular marginal folds (Reznik et al., 1978) that control movement of ingesta from esophagus to duodenum. An embryological study has shown that the forestomach is gastric in origin and not an esophageal derivative (Vorontsov, 1979). The structure and function of the forestomach is similar to the rumen of herbivores (Takahashi and Tamate, 1976; Borer, 1985). Ingesta enter the forestomach from the esophagus and pass into the glandular stomach in 10 to 60 minutes (Ehle and Warner, 1978). Kunstyr (1974) noted that the concentration of microorganisms is higher in the forestomach than in the glandular region. Sakaguchi et al. (1981) demonstrated that the forestomach aids in the utilization of dietary urea.

The hamster cecum is a J-shaped structure with numerous lateral sacculations and has more volume than the stomach (Krueger and Rieschel, 1950; Magalhaes, 1968).



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Nutrient Requirements of Laboratory Animals: Fourth Revised Edition, 1995 5 Nutrient Requirements of the Hamster Taxonomically hamsters are classified as a subfamily, Cricetinae, with 7 genera and 18 species in the family Muridae (Musser and Carleton, 1993). They are distributed throughout the Palearctic zone of Eurasia (Anderson and Jones, 1984). Original habitats of laboratory hamsters included clay deserts, shrub-covered plains, forested steppes, and/or cultivated fields. Golden hamsters, Mesocricetus auratus, collected from a burrow 8 feet deep in a wheat field near Aleppo, Syria, were established as a colony of laboratory animals at the Microbiological Institute of Jerusalem in 1930. These animals were used to complete research on kala-azar delayed by the failure of Chinese hamsters to breed in captivity (Adler and Theodor, 1931). Adler took breeding pairs to Paris and London to establish colonies at research institutions there (Bruce and Hindle, 1934; Adler, 1948; Murphy, 1985). Breeding stock were distributed to investigators in India, Egypt, and the United States (Doull and Megrall, 1939; Poiley, 1950). Golden hamsters is the hamster species most frequently used in research (Hoffman et al., 1968; Siegel, 1985; Van Hoosier and McPherson, 1987), but very little is known about their nutritional requirements. In the past 20 years, 7 additional hamster species have been used as laboratory animals. Animals identified as strain MHH:EPH are maintained in Hannover, Germany (Reznik et al., 1978; Mohr and Ernst, 1987). Mouse-like Chinese striped hamsters, Cricetulus barabensis, are used in research in cytogenetics, diabetes, and toxicology (Calland et al., 1986; Diani and Gerritsen, 1987). The large guinea-pig-like European hamster, Cricetus cricetus, formerly considered a pest in agricultural areas, is now a model for research in carcinogenesis. Dwarf hamsters, Phodopus campbelli and P. sungorus, of southern and western Siberia, are used in research in cytogenetics, carcinogenesis, diabetes mellitus, obesity, photoperiod changes, and social behavior (Pogosianz and Sokova, 1967; Hoffmann, 1973; Daly, 1975; Gamperl et al., 1978; Hoffmann, 1978. Steinlechner et al., 1983; Wade and Bartness, 1984; Pond et al., 1987; Ruf et al., 1991). Turkish hamsters, Mesocricetus brandti, are used in hibernation, taxonomy, and cytogenetics research (Lyman and O'Brien, 1977; Lyman et al., 1981, 1983; Todd et al., 1972). Colonies of the Romanian hamster, Mesocricetus neutoni , were established in Bucharest and used for research in cytogenetics and taxonomy (Hamar and Schutowa, 1966; Murphy, 1977; Popescu and DiPaolo, 1980). The Armenian, or migratory hamster, Cricetulus migratorius , is used on a limited basis in cytogenetics and oncology research (Lavappa and Yerganian, 1970; Cantrell and Padovan, 1987) (see Table 5-1). BIOLOGICAL AND BEHAVIORAL CHARACTERISTICS Unlike simple-stomached rats, mice, and guinea pigs, hamsters, like voles, have a stomach that consists of two distinct compartments: a keratinized, nonglandular forestomach (cardiac) separated from a glandular region (pyloric) by sphincter-like muscular marginal folds (Reznik et al., 1978) that control movement of ingesta from esophagus to duodenum. An embryological study has shown that the forestomach is gastric in origin and not an esophageal derivative (Vorontsov, 1979). The structure and function of the forestomach is similar to the rumen of herbivores (Takahashi and Tamate, 1976; Borer, 1985). Ingesta enter the forestomach from the esophagus and pass into the glandular stomach in 10 to 60 minutes (Ehle and Warner, 1978). Kunstyr (1974) noted that the concentration of microorganisms is higher in the forestomach than in the glandular region. Sakaguchi et al. (1981) demonstrated that the forestomach aids in the utilization of dietary urea. The hamster cecum is a J-shaped structure with numerous lateral sacculations and has more volume than the stomach (Krueger and Rieschel, 1950; Magalhaes, 1968).

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Nutrient Requirements of Laboratory Animals: Fourth Revised Edition, 1995 TABLE 5-1 Names, Characteristics, and History of Laboratory Hamsters Genus and Species Common Name or Origin Size Adult Weight, g Origin of Collection Where Used Introduction Date Cricetus cricetus Common hamster Large 337–500a Germany Germany 1971b Mesocricetus auratus Golden or Syrian Medium 120–180c Aleppo, Syria Palestine 1930d Mesocricetus brandti Brandt's or Turkish Medium 137–258e Turkey, Asia Minor United States 1965e Mesocricetus newtoni Newton's or Romanian Medium 120f E. Romania Romania 1964f Cricetulus migratorius Migratory or Armenian Small 40–80g Armenia United States, E. Europe 1963h Cricetulus barabensis Striped-back or Chinese Small 28–40i Wild, collected in Northeast China Eurasia, United States 1919j Phodopus campbellik Campbell's or Siberian Dwarf 30–50g Tuva, S. Siberia Eurasia, N. America 1965l Phodopus sungorusk Dzungarian or Siberian Variable; depends on season 25–45g Omsk, W. Siberia Eurasia, N. America 1968m a Reznik et al. (1978). b Reznik-Schüller et al. (1974). c von Frisch (1990). d Adler and Theodor (1931). e Lyman and O'Brien (1977). f Murphy (1977). g Cantrell and Padovan (1987). h Yerganian (1977). i Yerganian (1958). j Hsieh (1919). k Wilson and Reeder (1993). l Pogosianz and Sokova (1967). m Hoffmann (1978). Hamsters, like guinea pigs and gerbils, eat regularly, at 2-hour intervals throughout the day (Anderson and Shettleworth, 1977; Borer et al., 1979). Wild hamsters gather and store grains and other food in underground burrows to ensure a constant source of food (Borer et al., 1979; Micheli and Malsbury, 1982; Carleton and Musser, 1984). Hamsters are adapted to running and digging and are active primarily during twilight and during the night. The golden hamster has a gestation period of 15 to 18 days. Members of the Mesocricetus species are solitary animals that live in separate burrows with one or two chambers and entrances and exits; males and females meet only for breeding (Murphy, 1985). REPRODUCTION AND DEVELOPMENT Developmental and reproductive indices for three species are given in Tables 5-2 and 5-3, respectively. Young hamsters weigh 2 to 4 g at birth (see Table 5-4; Poiley, 1972). Average litter size is 11 (Slater, 1972), ranging from 2 to 16 (Anderson and Shina, 1972). Newborn hamsters are fetal in appearance—hairless, eyes and ears closed, and legs underdeveloped (Balk and Slater, 1987). Incisors are erupted at birth and young animals begin to eat solid food within 7 to 10 days (Balk and Slater, 1987). Hamsters weigh 40 g when weaned at 21 days (Poiley, 1972). Male hamsters are sexually mature at 42 days old, but females can breed as early as 28 to 30 days old (Selle, 1945; Balk and Slater, 1987). Litters with the greatest average number of pups are obtained from females 8 to 10 weeks old and males 10 to 12 weeks old (Robens, 1968; Balk and Slater, 1987). EXAMPLES OF PURIFIED AND NATURAL-INGREDIENT DIETS Two examples of purified diets and one of a natural-ingredient diet are presented in Tables 5-5 A-C, 5-6 A-C, and 5-7 A-C. These diets supported growth that was equivalent to the highest rates reported in our review of the literature. The two purified diets supported growth rates of 1.6 to 2.0 g/day. The natural-ingredient diet was selected from three that supported a growth rate of 1.9 g/day; of the three, it was intermediate in complexity. WATER AND ENERGY Male and female golden hamsters consume, on average, 8.5 mL water/100 g BW/day; males consumed 5 mL water/100 g BW/day, while females consumed 14 mL water/100 g BW/day (Fitts and St. Dennis, 1981). Thompson (1971) recorded Chinese hamsters intake of water to be 11.4 mL/100 g BW/day for males and 12.9 mL/100 g BW/day for females. Water intake for golden hamsters was found to be 4.5 mL/100 g BW/day in males and 13.6 mL/100 g BW/day for females. Little definitive work has been done on the energy requirement of the hamster, and few research studies include data on energy utilization. When fed a cereal-based diet containing 14.95 percent neutral detergent fiber (NDF) and 5.6 kcal gross energy (GE)/g diet (23.4 kJ/g diet), hamsters digested 45.2 percent of NDF and 81.5 percent of GE. Hamsters fed a 75 percent alfalfa meal diet that contained 40.6 percent NDF and 4.05 kcal GE/g diet (16.9 kJ/

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Nutrient Requirements of Laboratory Animals: Fourth Revised Edition, 1995 TABLE 5-2 Developmental Indices for Golden, Chinese, and Siberian Hamsters Variable Unit Amount Reference Goldena Birth weight g 2–3 Biven et al., 1987 Incisors present Day 1 Balk and Slater, 1987 Ears open Day 4–5 Balk and Slater, 1987 Eyes open Day 14–16 Balk and Slater, 1987 Solid food eaten Day 7–10 Balk and Slater, 1987 Weaning age Day 21   Weaning weight g 35–40 Harkness and Wagner, 1983 Mature weights g 95–150 female; 85–130 male Harkness and Wagner, 1983 Life span Year 1–3 Biven et al., 1987 Chineseb Birth weight g 1.5–2.5 Moore, 1965 Incisors present Day 1 Moore, 1965 Ears open Day 10–14 Moore, 1965 Eyes open Day 10–14 Moore, 1965 Solid food eaten Day 12 Smith, 1957 Weaning age Day 21 Calland et al., 1986 Weaning weights g 15–17 female; 16–17 male Avery, 1968 Mature weights g 25 female; 35 male Smith, 1957 Siberianc Birth weight g 1.8 Pogosianz and Sokova, 1967 Incisors erupt Day 0 Pogosianz and Sokova, 1967 Ears open Day 3–4 Pogosianz and Sokova, 1967 Eyes open Day 10 Pogosianz and Sokova, 1967 Solid food eaten Day 10 Pogosianz and Sokova, 1967 Weaning age Day 16–20 Pogosianz and Sokova, 1967 Weaning weight g 23 Pogosianz and Sokova, 1967 Mature weight g 30 female; 40–50 male Cantrell and Padovan, 1987 Life span Year 1–2 Cantrell and Padovan, 1987 a Golden hamster (Mesocricetus auratus). b Chinese hamster (Cricetulus barabensis) previously (C. griseus). c Campbell's, Djungarian, or Siberian hamster (Phodopus campbelli) (Pogosianz and Sokova, 1967); Siberian Dwarf, Djungarian, or Dzungrian hamster (Phodopus sungorus) (Cantrell and Padovan, 1987). g diet) digested the NDF and GE to the extent of 33.4 percent and 50.2 percent, respectively (Ehle and Warner, 1978). Arrington et al. (1966) reported that hamsters fed purified diets containing 12 and 16 percent casein had a total GE intake of 27 to 29 kcal/day (113 to 121 kJ/day) and gained 40 to 100 g over a 42-day period. Smaller hamsters (45 g) consumed 58 kcal/100 g BW/day (243 kJ/100 g BW/day), while larger hamsters (90 g) consumed 28 kcal/100 g BW/day (117 kJ/100 g BW/day). For a summary of energy balance in golden hamsters, see Borer (1985). LIPIDS An optimal concentration of dietary lipid has not been established for the hamster, although they seem to thrive on diets containing 4 to 20 percent fat (w/w). Knapka and Judge (1974) fed weanling (21 days old) male and female golden hamsters natural-ingredient pelleted diets containing 3.1, 5.0, 7.3, or 9.2 percent crude fat for 35 days. The feed:gain ratio decreased with increasing concentrations of dietary lipid. A decrease in feed intake was not associated with increased concentration of energy in the diet. Mortality was 1.0 percent when the diet contained 3.1 and 5.0 percent lipid. Higher mortality, but greater weight gain, occurred with the higher fat diets. The authors concluded that the lipid requirement for maximal growth of hamsters is slightly higher than 5 percent, but maximal growth should not be the only criterion used to determine the optimal concentration of lipid supplementation. Hamsters were maintained for a year or longer on starch gel diets (e.g., Table 5-6A) that contained up to 20 percent fat, with no mortality attributed to the fat load. However, feeding hamsters this diet for maintenance at 10 to 12 g/day [about 25 kcal ME/100 g BW/day (105 kJ ME/100 g BW/day)]

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Nutrient Requirements of Laboratory Animals: Fourth Revised Edition, 1995 TABLE 5-3 Reproductive Indices for Golden, Chinese, and Siberian Hamsters Variable Unit Amount Reference Goldena Breeding age Week 8–10, female; Balk and Slater, 1987     10–12, male Balk and Slater, 1987 minimum, female Day 28 Selle, 1945 Estrous cycle Day 4 Balk and Slater, 1987 Gestation Day 15.5 Balk and Slater, 1987 Litter size   4–16 Slater, 1972 average   11   Litters/lifetime   4–6 Balk and Slater, 1987 Reproductive life Month 10 Balk and Slater, 1987 Chineseb Breeding age Week 8–12, female; Calland et al., 1986   Week 32–48, male Calland et al., 1986 minimum, female Day 41 Moore, 1965 Estrous cycle Day 4 Moore, 1965 Gestation Day 19–21 Avery, 1968 Litter size   1–11 Calland et al., 1986 average   6.1 Calland et al., 1986 Litters/lifetime   5 Parkening, 1982 Reproductive life Month 16 ± 0.5 Parkening, 1982 Siberianc Breeding age Day 35–40 male 16 hours/light/day Cantrell and Padovan, 1987   Day 150 male 8 hours/light/day Cantrell and Padovan, 1987 Estrous cycle Day 4 Iakovenko, 1974 Gestation Day 18 Daly, 1975 Litter size   1–9 Pogosianz and Sokova, 1967 average   4–6 Pogosianz and Sokova, 1967 Litters/lifetime   12 Pogosianz and Sokova, 1967 Reproductive life Month 12 Pogosianz and Sokova, 1967 a Golden hamster (Mesocricetus auratus). b Chinese hamster (Cricetulus griseus and/or C. barabensis). c Siberian Dwarf, Djungarian, or Dzungrian hamster (Phodopus sungorus ) (Pogosianz and Sokova, 1967; Cantrell and Padovan, 1987); Campbell's, Djungarian, or Siberian hamster (Phodopus campbelli) (Pogosianz and Sokova, 1967; Iakovenko, 1974). tends to lessen obesity and the hypertriglyceridemia associated with allowing the animals free access to food (Hayes et al., 1993; K. C. Hayes, Brandeis University, personal communication, 1994) Signs of Lipid Deficiency A review by Holman (1968) reported that weanling hamsters fed a fat-free diet showed a slow rate of growth, had pale kidneys, and developed ulcers at the mucocutaneous junction of the anus. ESSENTIAL FATTY ACIDS n-6 Fatty Acids The requirement for n-6 fatty acids has not been determined for hamsters, but a deficiency has been demonstrated. Christensen and Dam (1952) found that feeding weanling hamsters a fat-free diet resulted in loss of hair, scaly skin, and development of a profuse secretion of cerumen (ear-wax)—a light-yellow, cholesterol-containing material. Signs of n-6 deficiency may be decreased by feeding hamsters a diet with 10 percent lard or a dietary supplement of 28 mg linoleic acid/day. n-3 Fatty Acids No studies are available on the distribution of n-3 fatty acids in hamster tissues or on the development of n-3 fatty acid deficiency. Three studies (Cunnane et al., 1985, 1986, 1987) were conducted in which ethanol was fed to hamsters with subsequent increases in n-9 fatty acids and decreases in the n-6 and n-3 fatty acids in liver triglycerides and phospholipids. Dietary treatments supplying a range of n-6:n-3 ratios were, predictably, found to influence tissue fatty acid composition in ethanol-fed and control hamsters.

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Nutrient Requirements of Laboratory Animals: Fourth Revised Edition, 1995 TABLE 5-4 Growth of Golden Hamster Outbred Cr:RGH (SYR)     Weight, g   Sex Age, days Average Range Male 1 2.9 2.0–4.0 Male 7 6.9 5.0–13.9 Male 14 18.3 11.5–33.2 Male 21 40.0 29.2–51.0 Male 28 48.6 32.0–70.8 Male 42 86.1 68.5–99.9 Male 56 91.5 85.6–97.5 Male 70 99.4 91.8–107.1 Male 84 103.9 98.9–109.2 Male 112 121.9 112.7–131.3 Male 140 131.8 116.6–145.6 Male 168 140.5 127.8–142.4 Female 1 3.0 2.0–3.8 Female 7 7.8 5.5–13.7 Female 14 17.4 11.0–31.6 Female 21 40.3 29.5–50.0 Female 28 44.1 31.0–68.5 Female 42 93.0 86.5–102.2 Female 56 94.5 85.3–104.1 Female 70 103.2 92.0–114.5 Female 84 114.9 103.3–126.5 Female 112 135.9 125.3–146.5 Female 140 149.6 147.0–159.8 Female 168 157.8 149.6–166.5 NOTE: In 10 of the 12 age groups, the females are larger than the males. SOURCE: Poiley (1972). TABLE 5-5A Rutten and de Groot Purified Diet for Hamsters Ingredient Amount, g/kg diet Casein 200.0a Wheat starch 635.0b Corn oil 50.0c Cellulose 50.0d Mineral mix 35.0e Vitamin mix 10.0f CaHPO4 15.0 DL-Methionine 3.0 Choline bitartrate 2.0 a Acid-precipitated, containing: protein 89.1% (N × 6.38); moisture 8.9%; ash 4.67%; pH of a 10% aqueous suspension, 4.5. b Ten percent of native wheat starch was replaced by pregelatinized wheat starch to provide pellets of suitable quality for feeding hamsters. c No antioxidants were added. d Dicacel, highly purified and bleached fibrous powder, consisting of 87–90% pure α-cellulose; average length of fibers ≈44 µm; water, 4%; ash, 0.12–0.15%; and lignin, 0.04%. e See Table 5-5B. Rutten and de Groot mineral mix based on AIN-76A. f See Table 5-5C. Rutten and de Groot vitamin mix based on AIN-76A. SOURCE: Rutten and de Groot (1992). TABLE 5-5B Rutten and de Groot Mineral Mix     Amount, g/kg   Compound Formula Mix Dieta Salt NaCl 110.0 1.51 Salt NaCl   2.34 Potassium citrate K3C6H5O·H2O 394.0 5.29 Potassium sulfate K2SO4 51.8 0.81 Potassium sulfate K2SO4   0.33 Magnesium oxide MgO 28.4 0.63 Manganese carbonate MnCO3·H2O 3.5 0.051 Ferric citrate FeC6H5O7·5H2O 24.0 0.173 Zinc carbonate ZnO·2CO3·4H2O 1.6 0.031 Cupric carbonate, basic CuCO3(OH)2·H2O 0.3 0.004 Potassium iodate KIO3 0.08 0.002 Sodium selenite Na2SeO3·5H2O 0.01 0.0001 Chromic potassium sulfate CrK(SO4)·12H2O 0.55 0.0025 Sodium fluoride NaF 0.063 0.001 Cobaltous chloride CoCl2·6H2O 0.127 0.002 Sucrose powder   385.57   a Amount of element (boldface element in the formula) provided when 35 g of mix is added per kg diet. SOURCE: Rutten and de Groot (1992). CARBOHYDRATES In hamsters fed diets containing 65 percent lactose or fructose, a mortality rate of 22 percent was observed, but only a 6 percent mortality rate was observed for hamsters fed 71 percent glucose, and 3 percent mortality with 62 percent sucrose (Gustafson et al., 1955). Salley and Bryson (1957) reduced mortality by decreasing the sugar to 54 percent and substituting either cornstarch or cellulose. TABLE 5-5C Rutten and de Groot Vitamin Mix   Amount, per kg   Compound Mix Dieta Retinyl palmitate/acetate 400,000.0 IU 4,000.0 IU Cholecalciferol 248,000.0 IU 2,480.0 IU All-rac-α-tocopheryl acetate 5,000.0 IU 50.0 IU Menadione Na-bisulfite 0.4 g 0.004 g Thiamin-HCl 2.0 g 0.020 g Riboflavin 1.5 g 0.015 g Pyridoxine-HCl 0.7 g 0.007 g Nicotinic acid 9.0 g 0.090 g Ca-d(+)-pantothenate 4.0 g 0.040 g Folic acid 0.2 g 0.002 g d(+)-Biotin 0.06 g 0.0006 g B12 0.005 g 0.00005 g Myo-inositol 10.0 g 0.1 g Sucrose powder to 1 kg     a Amount of vitamin provided as IU or g when 10 g of mix is added per kg of diet. SOURCE: Rutten and de Groot (1992).

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Nutrient Requirements of Laboratory Animals: Fourth Revised Edition, 1995 TABLE 5-6A Hayes Purified Diet for Hamsters Ingredient Amount, g/kg diet Casein 200.0 Glucose 170.0 Cornstarch (or rice flour) 366.0 Corn oil 50.0 Cellulose 100.0 Wheat bran 50.0 Mineral mixa 46.0 Vitamin mixb 12.0 Choline dihydrogen citratec 6.0 NOTE: Diets were fed as gel blocks, prepared by withholding from the formulation 60 g/kg of either rice flour or cornstarch and premixing it with 800 mL of water slowly heated to simmering to form a slurry that was then added to the remaining ingredients while mixing. a Hayes Mineral Mix (Hayes et al., 1989). See Table 5-6B for composition of mix. b Hayes-Cathcart Vitamin Mix. Mix modified based on K.C. Hayes, Brandeis University, personal communication (1993). See Table 5-6C for contents of mix. c Used in place of choline chloride for increased stability. SOURCE: Hayes et al. (1989). TABLE 5-6B Hayes Mineral Mix     Amount, g/kg   Compound Formula Mix Dieta Calcium carbonate CaCO3 290.4849 5.35 Calcium phosphate CaHPO4·2H2O 72.5970 0.78 Calcium phosphate CaHPO4·2H2O   0.60 Potassium phosphate K2HPO4 314.2049 6.49 Potassium phosphate K2HPO4   2.57 Magnesium sulfate MgSO4·7H2O 98.732 0.45 Magnesium sulfate MgSO4·7H2O   0.59 Sodium chloride NaCl 162.3664 2.94 Sodium chloride NaCl   4.53 Magnesium oxide MgO 32.0395 0.87 Ferric citrate FeC6H6O7·5H2O 27.0000 0.205 Potassium iodide KI 0.0774 0.0008 Potassium iodide KI   0.0027 Manganese sulfate MnSO4H2O 1.2211 0.0183 Manganese sulfate MnSO4H2O   0.0106 Zinc chloride ZnCl2 0.9149 0.0202 Cupric sulfate CuSO4·5H2O 0.2901 0.0034 Chromic acetate Cr(C2H3O2)3 0.0443 0.00046 Sodium selenite Na2SeO3 0.0043 0.00009 Sodium fluoride NaF 0.0232 0.00046 a Amount of element (boldface in formula) provided when 46 g mix is added per kg diet. SOURCE: Hayes et al. (1989), modified to correct published errors per K. C. Hayes, Brandeis University, personal communication, 1993. For correct version, see Hayes et al. (1993). TABLE 5-6C Hayes-Cathcart Vitamin Mix   Amount   Compound g/kg mix mg/kg dieta Retinyl palmitate (500,000 IU/g) 1.5 18.0 (9,000 IU) Cholecalciferol (400,000 IU/g) 0.1 1.2 (480 IU) All-rac-α-tocopheryl acetate (500 IU/g) 15.0 (7,500 IU) 180.0 (90 IU) Menadione 0.2 2.4 Myo-inositol 5.0 60.0 Niacin 3.0 36.0 Ca-pantothenate 1.6 19.0 Folic acid 0.200 2.4 Riboflavin 0.700 8.4 Thiamin 0.600 7.2 Pyridoxine-HCl 0.700 8.4 Biotin 0.020 0.24 Cyanocobalamin 0.001 0.012 Choline dihydrogen citrate 0.000 6,000.0 Dextrin 971.379   a When vitamin mix is added at 12 g/kg diet. SOURCE: Hayes et al. (1989). TABLE 5-7A Natural-Ingredient Diet for Hamsters Ingredients Amount, g/kg diet Alfalfa meal dehydrated (17% protein) 200.0 Corn, yellow dent ground grain 529.3 Soybean seed meal solv-extd (44% protein) 220.0 Dry beet molasses 7.0 Soybean oil 12.5 Salt 12.5 Dibasic calcium phosphate 18.0 Ground limestone 5.0 Trace mineral mixa 0.5 Vitamin mixb 0.8 Choline dihydrogen citratec 1.4 DL-Methionine 0.5 a Trace mineral mix formulated by E. A. Ulman provides minerals (g/kg): manganese (MnO2) 300; ferrous iron (FeSO4·7H2O) 570; zinc (ZnO) 97; copper (CuSo4),4; iodine (KIO3), 4.7; cobalt (CoCl2·6H2O), 3. For use, 0.5 g of mix is added per kg diet. b Vitamin mix designed by E. A. Ulman for use with cereal-based diets provides vitamins (g/kg) when added at 0.8 g/kg diet: vitamin A palmitate, 500,000 IU/g, 2.2; vitamin D, 250,000 IU/g, 4.7; vitamin E acetate, 500 IU/g, 52; menadione sodium bisulfite (62.5% menadione), 2.9; niacin, 19.5; Ca-pantothenate, 11.7; thiamin-HCl, 13.0; pyridoxin-HCl, 2.2; riboflavin, 4.4; folic acid, 2.9; biotin (1% in dextrose), 9.1; cyanocobalamin (1% in mannitol), 2.6; sucrose, 872.8 to give a total 1,000 g of mix. c More stable than choline chloride. SOURCE: Birt and Conrad (1981).

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Nutrient Requirements of Laboratory Animals: Fourth Revised Edition, 1995 TABLE 5-7B Trace Mineral Mix     Amount, g/kg   Compound Formula Mix Dieta Manganese dioxide MnO2 300.0 0.095 Ferrous sulfate FeSO4•7H2O 570.0 0.060 Zinc Oxide ZnO 97.0 0.039 Cupric sulfate CuSO4 24.0 0.0048 Potassium iodate KIO3 4.7 0.0014 Cobaltous chloride CoCl2•6H2O 4.3 0.0005 a Amount of element (boldface in formula) provided when 0.5 g mix is added per kg diet. SOURCE: Formulation by E. A. Ulman based on Birt and Conrad (1981). With purified, fiber-free diets containing 64 percent carbohydrate, cornstarch was superior to glucose or sucrose in supporting survival (Ershoff, 1956). Rice starch supported higher growth rates than lactose (Dam and Christensen, 1961). Rogers et al. (1974) obtained satisfactory growth in a long-term study when animals were fed a gel diet containing 40 percent cornstarch and 21.9 percent sucrose. Hayes et al. (1989) observed that ''wet tail" could be prevented by inclusion of rice flour, fiber, or lactose in gel diets. The implication is that diarrhea and "wet tail," commonly encountered in hamsters fed purified diets, results from an insufficient amount of complex carbohydrates (fiber, starch) reaching the large bowel flora. PROTEIN AND AMINO ACIDS Compared to a ruminant, fermentative digestion in the hamster is not sufficient to alter the pattern of dietary amino acids enough to improve growth of hamsters fed proteins such as wheat gluten (Banta et al., 1975). It seems that hamsters can make limited use of urea as a source of TABLE 5-7C Vitamin Mix   Amount   Compound g/kg mix per kg dieta Vitamin A palmitate 500,000 IU/kg 2.2 880.0 IU Vitamin D2 50,000 IU/g 4.7 188.0 IU Vitamin E acetate 500 IU/kg 52.0 20.8 IU Menadione sodium bisulfite 62.5% menadione 2.9 1.5 mg Nicotinic acid 19.5 15.6 mg Ca-pantothenate 11.7 9.4 mg Thiamin-HCl 13.0 10.4 mg Pyridoxine-HCl 2.2 1.8 mg Riboflavin 4.4 3.5 mg Folic acid 2.9 2.3 mg Biotin (1% in dextrose) 9.1 0.07 mg Cyanocobalamin (1% in mannitol) 2.6 0.02 mg Sucrose powder 872.8   a Value when mix is added at 0.8 g/kg diet. SOURCE: Formulation by E. A. Ulman based on Birt and Conrad (1981). This vitamin mix is designed to be used with cereal-based diets. dietary nitrogen (Matsumoto, 1955; Sakaguchi et al., 1981). However, fermentative digestion seems to suppress the anticipated response to supplementation of amino acids expected to improve growth (Arrington et al., 1966; Banta et al., 1975) and to decrease the toxicity of high dietary concentrations of L-phenylalanine (Horowitz and Waisman, 1966). GROWTH No studies to determine requirements for single amino acids or mixtures of amino acids were found. Studies that focused on protein requirements used variations in natural-ingredient diets or, in a few cases, used a single protein product at a series of dietary concentrations (Table 5-8). Protein requirements for growth reported here were ob- TABLE 5-8 Protein Requirements Strain Growth, g/day Protein Source Amounts, % Reference Syrian 1.8–2.0 Corn, soybean, casein 13.7 Arrington et al., 1979 Ufnl:(SYR)a         Golden 1.8–1.9 Multiple natural-ingredients 20 Banta et al., 1975 Syrian   Wayne Lab Blox 24 Birt and Conrad, 1981     Teklad 22 Birt and Conrad, 1981     Corn-soybean 18 Birt and Conrad, 1981   1.8–1.9 Corn-soybean-alfalfa 18 Birt and Conrad, 1981     Multiple natural-ingredients 20   Syrian 1.2 Multiple natural-ingredients 18 Feldman et al., 1982 Lak:LVG (Syr)a         Syrian 1.4 Casein 18 Arrington et al., 1966 Syrian 1.4 Casein 18 Horowitz and Waisman, 1966 NOTE: In all studies, both males and females were used. a Only groups identified as strains.

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Nutrient Requirements of Laboratory Animals: Fourth Revised Edition, 1995 tained from studies that used both male and female 3- to 4-week-old hamsters weighing approximately 40 g. Growth rate varied from 1 to 2 g/day in experiments lasting 3 to 5 weeks in most cases. Arrington et al. (1979) found that diets containing 13.7 percent crude protein from mixtures of corn, soybean, and casein would support gains of 1.8 to 2 g/day in both male and female golden Syrian hamsters. Banta et al. (1975) reported growth rates of 1.8 to 1.9 g/day during a 6-week period using a natural-ingredient diet containing 20 percent crude protein. Birt and Conrad (1981) compared two commercial natural-ingredient diets with three formulated diets of increasing complexity and found that diets that contained 18 to 22 percent crude protein would support gains of 1.8 to 1.9 g/day in both male and female hamsters over a period of 6 weeks. However, Feldman et al. (1982) obtained maximum gains of only 1.2 g/day even when the natural-ingredient diets contained up to 24 percent crude protein. A natural-ingredient diet containing 18 percent crude protein should support growth rates approaching 2 g/day in weanling hamsters. Addition of semipurified proteins to natural-ingredient diets may reduce the amount of crude protein required to support the expected rates of growth. In studies using semipurified sources of protein, Arrington et al. (1966) found that diets containing 18 percent casein resulted in growth of 1.4 g/day over 6 weeks. Additional experiments with diets containing 16 percent casein or soy protein isolate supported gains of 1.4 to 1.5 g/day in experiments lasting 5 weeks (Arrington et al., 1966). In another study using casein at 9, 18, or 25 percent of the diet, Horowitz and Waisman (1966) found that maximum gain (1.4 g/day) was obtained with 18 percent casein. Variation in the age and size of hamsters used in the experiments reviewed make it difficult to determine whether lower rates of growth obtained when purified proteins were used is the result of the hamster or the diet. Addition of free amino acids to the diet to improve the dietary amino acid pattern and, hence, growth have not been successful. A form of encapsulated amino acid may be beneficial. REPRODUCTION Two studies focused on diet and reproduction in hamsters (Birt et al., 1982; Birt and Conrad, 1981). In one study (Birt and Conrad, 1981), the effects of five natural-ingredient diets containing from 18 to 24 percent crude protein (see, for example, Table 5-7A) were compared over three breeding cycles; reproductive performance of hamsters fed corn-soybean or corn-soybean-alfalfa meal diets containing 18 percent crude protein was equal to or exceeded that of hamsters fed either of two commercial natural-ingredient diets containing 22 to 24 percent crude protein. In the other study (Birt et al., 1982), which used lactalbumin at 20 and 40 percent of the diet as the sole source of protein, reproductive efficiency (pups weaned per mating) was 20 to 40 percent that of identical females fed the commercial diet. Although no recommendation is made for the amount of purified protein required to support reproduction, a natural-ingredient diet containing 18 percent crude protein is thought to meet the amino acid needs for reproduction in hamsters. MINERALS Given their widespread use as experimental animals, there is a remarkable paucity of information about the mineral requirements of hamsters. MACROMINERALS Calcium and Phosphorus Normal bone formation occurred in hamsters fed diets containing 6.0 g Ca/kg and 3.5 g P/kg. In the absence of vitamin D, rickets was produced in hamsters fed 4.7 g Ca/kg and 2.0 g P/kg diet (Jones, 1945). Old female hamsters fed diets containing 4.0 g P/kg and 3.0, 5.0, or 7.0 g Ca/kg were in positive calcium balance only at the two higher calcium intakes. Young animals (52 days old) retained calcium when fed 3.0, 5.0, and 7.0 g Ca/kg diet (Kane and McCay, 1947). Stralfors (1961) obtained a 54 percent decrease in the incidence of dental caries in hamsters when the calcium content of the diet was increased from 4.0 to 6.0 g Ca/kg. Sodium and Chloride Rowland and Fregly (1988) reported that hamsters, unlike rats, were reluctant to ingest NaCl either spontaneously or after treatment with several natriogenic stimuli that were effective in rats. Furthermore, they noted that variations in intake of NaCl solutions made hamsters extremely refractory to either decreases or increases in functional mineralocorticoid activity. TRACE MINERALS No studies were located that specifically addressed the dietary requirements of the hamster for iodine, molybdenum, and selenium or for iron. Iodine, Molybdenum, and Selenium Iodine, molybdenum, and selenium are trace elements essential for normal growth in laboratory animals. Birt et al.

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Nutrient Requirements of Laboratory Animals: Fourth Revised Edition, 1995 (1986) determined that male and female golden hamsters fed a diet containing 30 percent torula yeast as the protein source and 0.1 mg Se (as sodium selenite)/kg diet supported adequate growth, and 5 mg Se/kg is excessive. The iodine requirement may be met by 0.15 mg I/kg diet, and the molybdenum requirement by 0.10 mg Mo/kg diet. The selenium requirement may be met by 0.15 mg Se/kg diet for maintenance, 0.20 for growth and aging, and 0.40 for pregnancy and lactation. Signs of Iodine Deficiency Hamsters fed iodine-deficient diets (10 to 25 µg/kg) for several months developed enlarged thyroids when compared to controls fed adequate iodine (7.6 mg/kg) (Follis, 1959, 1962). Iron Signs of Iron Deficiency Chandler et al. (1988) reported that mild iron deficiency can be induced in adult males by feeding them a diet containing 10 mg Fe/kg for several weeks. Carpenter (1982) reported that feeding females a low-iron diet (3 mg Fe/kg) during pregnancy resulted in low maternal weight gain and a high frequency of prenatal mortality compared to controls. VITAMINS FAT-SOLUBLE VITAMINS Vitamin A The vitamin A requirement of golden hamsters seems to be only slightly greater than that of the rat. Hamsters fed a purified diet containing 2 mg retinyl palmitate/kg diet (3.8 µmol/kg) grew as well as animals fed a commercial natural-ingredient diet (Rogers et al., 1974). The hamsters had normal serum vitamin A concentrations and a very modest accumulation of vitamin A in the livers. Based on these studies the minimum amount of retinol that will maintain a slightly positive vitamin A balance is approximately 1.1 mg/kg diet (3.8 µmol/kg diet). Signs of Vitamin A Deficiency Omission of vitamin A from a 24 percent-casein diet resulted in deficiency signs in 6 to 7 weeks. Vitamin A-deficient animals developed abnormally and had coarse and sparse hair, xerophthalmia, and keratinized stratified tracheal lining (Salley and Bryson, 1957). Stomach ulcers formed in adult male hamsters fed a vitamin A-deficient diet for 7 months (Harada et al., 1982). Signs of Vitamin A Toxicity Hamsters fed a diet containing 400,000 IU vitamin A/kg (419 µmol/kg) developed liver pathology and died within 42 to 91 days. Animals fed 100,000 IU/kg diet (105 µmol/kg) and lower concentrations (4,000 and 600 IU/kg) showed no toxic effects (Beems et al., 1987). Vitamin D Overt signs of rickets did not appear within a 5-week period when hamsters were fed vitamin D-deficient diets that contained calcium and phosphorus in a ratio of 2:1 and calcium was included at 6 g/kg diet. Rickets may be induced in hamsters in the absence of vitamin D and when dietary calcium is 4 g/kg and phosphorus is 0.2 g/kg (Jones, 1945). No published reports on vitamin D deficiency or toxicity were found. Vitamin E In spite of several studies on vitamin E deficiency in hamsters and their use as an animal to bioassay compounds for vitamin E activity, data are not available to provide a good estimate of the vitamin E requirement of hamsters. Bieri and Evarts (1974) found that a diet containing 2.1 µmol/kg RRR-α-tocopheryl acetate/kg was adequate to prevent testicular degeneration in the rat. However, plasma creatine phosphokinase (CPK) concentrations were slightly higher at this concentration of intake. With higher dietary concentrations (6.3 µmol/kg), plasma CPK concentrations were normal (Bieri, 1972). Unfortunately the more sensitive criterion of vitamin E adequacy, such as the in vitro red blood cell hemolysis assay, has not been investigated in hamsters. In the rat 6.3 µmol RRR-α-tocopherol/kg diet may be adequate to prevent overt signs of vitamin E deficiency, but this concentration is quite likely not adequate for optimal performance. Therefore, 42 µmol RRR -α-tocopherol/kg diet (27 IU/kg), which is required by the rat, is probably a more realistic value. A few breeding colonies have reported a higher-than-expected incidence of spontaneous hemorrhagic necrosis, a fatal disease that affects the central nervous system of fetal hamsters. Keeler and Young (1979) found that a single intraperitoneal injection of 100 µmol of vitamin E on day 7 of gestation protects fetuses from this disease. The problem may arise from improper storage of diets, which leads to destruction of vitamin E. Signs of Vitamin E Deficiency The absence of vitamin E in their diet causes hamsters to develop testicular degeneration. Feeding a hamster 21 µmol RRR-α-tocopheryl acetate/day restores normal weight and testicular histology. In contrast, rats are unable to reverse vitamin E-induced testicular degeneration (Mason and Mauer, 1975). Vitamin E-deficient hamsters show decreased growth and muscular dystrophy, which can be alleviated by administering high

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Nutrient Requirements of Laboratory Animals: Fourth Revised Edition, 1995 concentrations of vitamin E (West and Mason, 1958). Weanling hamsters fed a vitamin E-deficient diet developed muscular degeneration and died within 2 weeks. Improvement occurred within 30 hours after a single dose of 1 mg α-tocopherol (Houchin, 1942). Vitamin K No studies to ascertain the hamster's requirement for vitamin K could be found; however, Rogers et al. (1974) reported that a diet containing 4 mg menadione/kg (23 µmol/kg diet) was adequate for growth—the hamsters presumably received a considerable amount of vitamin K activity from coprophagy. No bleeding problems were reported. The Tolworth HS (Welsh) Warfarin-resistant strain of rats requires 1.77 µmol phylloquinone/kg BW/day (Greaves and Ayers, 1973). The vitamin K requirement of the hamster may be similar. Based on the requirement of the Tolworth HS (Welsh) Warfarin-resistant rat, 25 µmol phylloquinone/kg diet (11 mg/kg) should be a safe and adequate intake for hamsters. Signs of Vitamin K Deficiency Adult male hamsters fed a vitamin K-deficient diet and housed in coprophagy-preventive cages showed a drop in prothrombin concentrations to 11 percent of control concentrations within 11 days. Treatment with chloro-K (2-chloro-3-phytyl-1,4-naphthoquinone), a vitamin K antagonist, at 1 to 5 mg chloro-K/kg BW decreased plasma prothrombin to 17 to 20 percent of control values. Hamsters are Warfarin resistant and require a large amount to reduce prothrombin production (Shah and Suttie, 1975). WATER-SOLUBLE VITAMINS Biotin Satisfactory growth of hamsters has been obtained with diets containing 0.82 µmol biotin/kg (Cohen et al., 1971) and 2.5 µmol biotin/kg (Rogers et al., 1974). A dietary concentration of 0.2 mg/kg (0.82 µmol/kg) seems to be a safe and adequate amount of biotin for hamsters. Under normal conditions golden hamsters do not require dietary biotin (Granados, 1968). Apparently, the biotin obtained through coprophagy is sufficient to meet the requirement. Signs of Biotin Deficiency A biotin deficiency is induced by feeding hamsters a diet containing both raw egg white and sulfaguanidine. Biotin-deficient animals developed dull rough coats, encrusted eyes, depigmented hair, and jerky movements. Daily injections of 16 nmol biotin (3.9 µg), equivalent to 0.66 mg/kg diet (2.7 µmol/kg), reversed deficiency signs within 4 to 6 weeks (Rauch and Nuting, 1958). Ten adult female hamsters fed a purified diet containing 5.0 mg biotin/kg diet produced 118 normal live fetuses; but 11 animals fed a biotin-deficient diet had 20 live fetuses (Watanabe and Endo, 1989). Choline Hamsters fed a peanut meal diet deficient in choline developed poor appetite, reduced growth, and fatty livers (Handler and Bernheim, 1949). Investigators who fed hamsters diets containing more than 200 g casein/kg did not observe a requirement for choline (Hamilton and Hogan, 1944). Purified diets on which hamsters have achieved satisfactory growth have contained 14 µmol choline chloride/kg diet (Rogers et al., 1974) or 7.1 µmol choline bitartrate/kg diet (Cohen et al., 1971). Under most circumstances 1.8 g choline bitartrate/kg diet should provide a safe and adequate intake of choline. Folates Golden hamsters fed 2 mg folic acid/kg diet (Cohen et al., 1971) do not develop folate deficiency. Folate deficiency does develop when golden hamsters are fed a purified diet containing 60 percent sucrose or 1 percent sulfonamide with either cornstarch or sucrose as the carbohydrate. Hamsters resemble guinea pigs rather than rats in that a folate deficiency can be produced without the use of sulfonamide. Signs of Folate Deficiency In hamsters given a folate-deficient diet, liver folates are decreased to 10 percent of control and blood PCV and hemoglobin values are decreased 20 percent. In the deficient animal, increases are seen in urinary excretion of formiminoglutamic acid and aminoimidazolecarboxamide (Cohen et al., 1971). Myo-inositol Granados (1968) has stated that hamsters do not require Myo-inositol for normal growth. Hamilton and Hogan (1944) also demonstrated that myo-inositol is not required for growth but is necessary for reproduction. Niacin Young hamsters fed diets containing 20 percent casein do not require dietary niacin for growth (Hamilton and Hogan, 1944; Granados, 1951). Niacin is necessary, however, for normal reproduction and adequate litter size (Hamilton and Hogan, 1944). Signs of Niacin Deficiency Hamsters fed a niacin-free, purified diet developed rough and denuded hair, suffered loss of weight and death. Hair quality and weight gain

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Nutrient Requirements of Laboratory Animals: Fourth Revised Edition, 1995 improved with daily administration of 100 µg niacin (Routh and Houchin, 1942). Niacin supplementation does not improve growth of hamsters fed 20 percent casein (Hamilton and Hogan, 1944; Granados, 1951). However, niacin probably will be required in diets with low concentrations of protein or diets in which tryptophan is first limiting. Animals fed purified diets containing 81 µmol niacin/kg achieved satisfactory growth rates over 11 weeks (Cohen et al., 1971). Pantothenic Acid Routh and Houchin (1942), Hamilton and Hogan (1944), Granados (1951), and Cohen et al. (1963) identified pantothenic acid as a nutrient required for normal growth. Nevertheless, the requirement for this vitamin has not been quantified in the hamster. Concentrations of dietary Ca-pantothenate from 21 µmol/kg (Hamilton and Hogan, 1944) to 84 µmol/kg diet (Rogers et al., 1974) have been used in purified diets. None of the investigators indicated whether the type used was Ca-d-pantothenate or Ca-dl-pantothenate. Thus, 21 µmol Ca-d-pantothenate/kg diet (10 mg/kg diet) seems to be a safe and adequate concentration of pantothenic acid activity. Signs of Pantothenic Acid Deficiency In the absence of pantothenic acid, hamsters lost weight, developed a red encrustation around the mouth, and died in 20 days. Daily injections of 15 µg Ca-pantothenate supported maintenance, but larger doses were needed for growth (Routh and Houchin, 1942). Vitamin B6 Male weanling hamsters fed a pyridoxine-deficient diet for 2 to 3 weeks decreased their food and water intake and stopped growing. No quantitative requirement for vitamin B6 can be set at this time. Signs of Vitamin B6 Deficiency In addition to weight loss, the hair of vitamin B6-deficient hamsters was unkempt, and crusted lesions were occasionally observed on lips and mouth. Increased xanthurenic acid was found in urine. Atrophy of lymphoid tissue, particularly in the thymus, is an outstanding pathological change (Schwartzman and Strauss, 1949). In the absence of vitamin B6, hamsters lose weight and develop an acrodynia-like condition around the mouth in less than 2 weeks. A daily dose of 3 µg pyridoxine cured the dermatitis and produced moderate growth (Rouch and Houchin, 1942). Riboflavin Riboflavin is required for normal growth and development of hamsters, but no quantitative requirement has been established (Hamilton and Hogan, 1944; Granados, 1968). Deficiency signs did not occur in animals fed 20 mg/kg diet (Smith and Reynolds, 1961). Riboflavin depletion (measured by erythrocyte glutathione reductase) was produced in hamsters fed two concentrations of riboflavin—0.5 and 1.5 mg/kcal—in a liquid diet. Decreased growth was observed at the lower dose (Kim and Roe, 1985). Thus, a diet containing 15 mg/kg should be sufficient to support normal growth in hamsters. Signs of Riboflavin Deficiency In the absence of dietary riboflavin, hamsters reduced their food and water intake, became inactive, showed stunted growth, and developed dull coats (Smith and Reynolds, 1961). Thiamin Thiamin is necessary for normal growth, but the specific requirement has not been established. Satisfactory growth has been obtained with purified diets containing 20 mg thiamin/kg diet (Arrington et al., 1966). Signs of Thiamin Deficiency Hamsters fed 4 mg thiamin/kg diet developed a chronic deficiency (Salley et al., 1962). Hamsters fed a thiamin-deficient diet developed polyneuritis in 12 days. Oral administration of 3 µg thiamin/day reversed these signs in 2 days (Routh and Houchin, 1942). Vitamin B12 Early reports concluded that vitamin B12 is not required for normal growth of golden hamsters (Scheid et al., 1950; Granados, 1951). Hamsters fed a diet high in soybean protein and cornstarch showed a mild vitamin B12 deficiency identified by the presence of distinctive metabolites (methylmalonic acid and formiminoglutamic acid) in urine (Cohen et al., 1967). Metabolic changes in deficient animals were corrected by feeding them a diet containing 10 µg vitamin B12/kg diet. The inclusion of inorganic cobalt (5 mg/kg diet) in the diet reversed deficiency changes and increased tissue storage of vitamin B12 (Tseng et al., 1976). POTENTIALLY BENEFICIAL DIETARY CONSTITUENTS FIBER Fiber-free diets containing high concentrations of purified sugars result in high mortality (Salley and Bryson, 1957). The substitution of cornstarch for glucose and sucrose or addition of 12 to 20 percent alfalfa to diets increased survival (Ershoff, 1956). Basal diets containing

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Nutrient Requirements of Laboratory Animals: Fourth Revised Edition, 1995 starch or lactose may not require fiber additions because these ingredients support favorable microflora in the colon (Snog-Kjaer et al., 1963; Hayes et al., 1989). Microoorganisms present in the cecum and colon seem to be capable of degrading fiber sources (Banta et al., 1975; Vorontsov, 1979). Hamsters, like other rodents, practice coprophagy (von Frisch, 1990). ASCORBIC ACID Early work suggested that golden hamsters do not require a dietary source of ascorbic acid (Clausen and Clark, 1943). Male hamsters fed a purified diet supplemented with 4.0 mg ascorbic acid/g diet gained 1.07 g/day with a food intake of 3.5 g/day. Growth curves overlapped; curves for supplemented animals were slightly above those of controls, which were fed diets containing no ascorbic acid. None of the animals weighed more than 96 g after 120 days. Poiley (1972) reported average male weight as 122 g at 112 days (Table 5-4). Forty female hamsters fed a diet scorbutic for guinea pigs were given daily supplements of ascorbic acid in water by pipette according to the weight/dose chart of Dann and Cowgill (1935). Controls received this diet and an equal amount of water. At 70 days average weights were 105.6 g with a 0.3 mg dose, 99.4 g with a 0.65 mg dose, and 91.0 g with 0.9 mg dose per 100-g animal per day. Controls fed water and a diet scorbutic for guinea pigs averaged 88.5 g. Animals fed water and a diet supplemented with lettuce averaged 92.3 g (Hovde, 1950). Poiley (1972) reported, on average, females weighed 103 g at 70 days. REFERENCES Adler, S. 1948. Origin of the golden hamster Cricetus auratus as a laboratory animal. Nature 162:256–257. Adler, S., and O. Theodor. 1931. Investigations on Mediterranean kala azar. Proc. R. Soc. Biol. 108:453–463. Anderson, M. C., and S. T. Shettleworth. 1977. Behavior adaptation to fixed-interval and fixed-time food delivery in golden hamster. J. Exp. Anal. 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