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Intake by Puppies
OLAV T. OFTEDAL1
Milk composition and yield vary greatly age or less are reported to consume the
among diverse mammalian species (1-3). equivalent of 10-14% of body weight per day
Estimation of the nutrient requirements of (7, 9). The effects of maternal nutrition (9,
both mother and suckling young requires 10), maternal size (2), breed (6), and litter size
quantitative information on lactation per- and mass on lactation performance in dogs
formance. Dogs are known to produce a need clarification.
rather concentrated milk containing 21-26% The following study was undertaken to
total solids, 8-12% fat and 7-10% protein measure milk composition and milk yield at
(4-7), although lower levels of fat and protein peak lactation in well-nourished dogs of the
have recently been reported (8). Little in- beagle breed. This study is part of a larger
formation is available on milk yields in dogs, project in which lactation performance is
Several litters of various breeds have been compared among several species, and nutri-
studied by weighing puppies before and after ent intakes of suckling young are related to
suckling (6, 7). A German shepherd was esti- body size, growth rates and estimated re-
mated to produce 1.7 kg milk per day at the quirements (3, 11). Milk production was
lactation peak at 3 weeks postpartum, where-
aS bitches Of Smaller breeds produced leSS © 1984 American Institute of Nutrition. Received for publication
milk but were only studied in the first 9 days 27,]une1983M
,„. ^ ... , . J - Current address: Department of Zoological Research, National
(b). bUCkling pUppieS 4 WeekS Or Zoological Park, Smithsonian Institution, Washington, DC 20008.
803
804 OFTEDAL
measured from the dilution and turnover of tocin (5 IU) was administered by intramus
deuterium oxide (D2O) administered to cular injection, and one or two teats evacu
puppies. Hydrogen isotopes have been ated as completely as possible by gentle
shown to yield valid estimates of milk manual expression. An average of 21 ml
production if corrections are made for ( ±6.3 SD) was obtained in 10-15 minutes.
changes in body water pool size and for Samples were frozen in sealed vials until
isotope recycling via maternal milk (11-15). analyzed.
Peak lactation was assumed to occur in week Milk samples were thawed quickly, ho
3 or 4 postpartum since puppies do not mogenized in a Potter-Elvehjem tissue
initiate feeding on semisolid food until the grinder and subsampled. The weekly sam
emergence of deciduous dentition at 21 to 35 ples were assayed in duplicate for major
days postpartum (16-18). Milk alone will constituents. Total solids were determined
support normal growth up to 4 weeks post by oven drying, total nitrogen (TN) and
partum; thereafter, withholding of supple nonprotein nitrogen (NPN) by a Kjeldahl
mental food may result in a reduced growth procedure, fat by the Roese-Gottlieb method
rate (19). (21) and sugar by the phenol-sulfuric acid
colorimetrie method, as previously described
grating infrared spectrophotometer (Model ance (ANOVA)]. Sugar content did differ
521, Perkin-Elmer Corp., Norwalk, CT). among sampling times (P < 0.05, ANOVA),
Assayed deuterium levels were corrected the mean value rising from 3.47% at 7 to 9
for body weight changes in computations of days postpartum to 4.13% at 29 to 30 days
fractional turnover rate (k) and body water postpartum. The mean values for all sam
fraction (F)(ll). Isotope recycling via mater pling times were: 22.7 ±0.41% total solids,
nal ingestion of the excreta of suckling 9.47 ±0.386% fat, 7.53 ±0.123% protein,
young, followed by transfer of isotope in 3.81 ±0.079% sugar and 146 ±3.6 kcal
milk water from mother to young (12), gross energy per 100 g. If converted to a dry
necessitated an additional correction. The matter basis dog milk was found to contain
accumulation of deuterium in an unin- 41.4 ±0.87% fat, 33.4 ±0.60% protein,
jected, control puppy in each litter was 17.0 ±0.049% sugar and 641 ±3.9 kcal/
monitored. On the assumption that these 100 g dry matter. Fat, protein and sugar
levels are representative of recycled isotope provided 58.7 ±0.86%, 30.5 ±0.66% and
in littermates, the deuterium levels in con 10.5 ±0.33% of total gross energy, respec
trol puppies were subtracted from the tively. There were no significant differences
TABLE 1
Composition of dog milk1'*
blood sampling in the first and second study Milk intakes of puppies were compared
periods, respectively. Correction for isotope among litters and between the two post
recycling was therefore warranted. natal ages by two-way ANOVA. Whether
Fractional turnover rate of body water (k) expressed as a daily amount, as a percentage
declined from 0.168 ±0.0036 per day for of body weight per day, or per gram body
the first study period to 0.152 ±0.0034 per weight gain, mükintake was significantly
day for the second (P < 0.001, paired i-test). influenced by both litter and age effects
By contrast body water fraction (F) did not (table 3). Although the absolute amount of
differ significantly (P > 0.05, paired f-test) milk consumed per day at 26 days (175
between the two periods (table 2). Calcu ±5.3 g) was greater (P < 0.01) than that at
lated water losses, water gains and water 19 days (160 ±5.4 g), this amount repre
intakes at 19 and 26 days postpartum are sented a smaller percentage of body weight
presented in table 2. Water intake was (14.6% at 26 days vs. 17.0% at 19 days, P
equivalent to 15.1 and 13.9% of body weight < 0.001). Since the same estimate of growth
at 19 and 26 days, respectively. rate was used for both age categories, milk
Water intake derives from both preformed intake per gram body weight gain was of
milk water and metabolic water from the course greater at 26 days (table 3). The
TABLE 3
Milk intakes of suckling puppies'
postpartum18.714.816.717.317.34
days
BR82 203152154156147160 4.51 1015
BR50 4.02 760
CA46 4.13 924
CD 18 4.54 1092
CA45 4.55 1029
All litters ± 0.3026 17.0 ±
5.221166178165159175 4.37 ±0.074 964 ±57.6
postpartum15.712.815.114.414.83
ilnu\
total solids, 9.5% fat, 7.5% protein and determination of fat depends on the degree
3.8% sugar. Mean values from prior studies of unsaturation of the lipids (34). Binding of
on dog milk are tabulated for comparison Coomassie brilliant blue G250 dye to pro
(table 4). This list includes 19th century tein is likewise a function of the amino acid
results of questionable analytical accuracy composition of the protein (35, 36). These
as well as studies involving only a few sam methods are valid only if standardized to the
ples from one or two dogs. Samples collected particular mix of lipid and protein constitu
very early or late in lactation have been ents found in dog milk. It appears that this
excluded as not representative of established was not done
lactation. Despite variation in sampling and The variation among the remaining
analytical procedures, most reports fall studies may be a function of sampling or
within the ranges of 21-26% total solids, analytical bias or may represent real differ
8-12% fat, 7-10% protein and 3-4% sugar ences among dogs. Dog breeds vary tre
(table 4). The results reported herein are mendously in body size and conformation,
consistent with these values. By contrast the but no correlation to the gross composition
recent data of Lönnerdal and colleagues (8) of milk could be determined by Russe (6)
indicate much lower fat (4.8%) and protein who studied breeds ranging in size from
(5.2%) levels in beagle milk collected 11-40 dachshunds to Saint Bernards. Beagle milk
days postpartum. This discrepancy may collected by Luick and colleagues (5) con
stem from inappropriate application of tained more total solids, fat and protein, but
rapid spectrophotometric methods. Color less sugar (table 4) than was found in the
development in the sulfuric acid-phosphoric present study. By contrast the recent data of
acid-vanillin reaction employed in the Mundt and colleagues (7) are very similar to
808 OFTEDAL
TABLE 4
Published data on the composition of dog milk at midlactation
Females Days after No. of Total
Source milked birth samples solids Fat Protein Sugar
(26)Tolmatscheff
Ssubotin 1866
(27)Abderhalden 1867
(28)Abderhalden1898
(29)Dijkstra 1899
(30)Grimmer
1910
(31)Daggs 1915
(10)Deniges
1931
(32)Anderson
1935
(4)Luick et al. 1940
(5)Russe
et al. 1960
(6)Lauer
1961
(33)Mundt
et al. 1969
the present results although four breeds as The NPN content of dog milk averaged
well as mongrels were included. Breed dif 0.054%. The data of Grimmer (31) and
ferences in the composition of dog milk Russe (6) indicate mean NPN values of 0.068
must be minor if they exist at all. and 0.112%, respectively. In the present
An elevation in total solids and protein at study NPN accounted for only 4.4% of total
both the beginning and the end of lactation nitrogen, as compared to 5.7 (31) and 9.3%
has been noted in prior reports (4, 6, 31). In (6). Protein estimates based on total nitro
the present study there were no substantial gen (e.g., see refs. 4, 5, 7, 33) will over
changes in milk composition from 7 to 37 estimate true protein by an amount equal to
days postpartum, a week after supplemental NPN x 6.38, i.e., by about 0.3 to 0.7
feeding of puppies commenced. Marked percentage points.
compositional changes are apparently asso In 100 g milk, 1.88 ±0.062 g of solids
ciated with mammary involution at about was not accounted for by the summation of
39—49days (4, 6). Weaning is of course fat, protein and sugar. Part of the residual is
influenced by feeding and management due to ash and part to NPN constituents. If
practices, and perhaps by breed as well (6). published data on the ash content of dog
Lonnerdal et al. (8) have reported a rise in milk in the period of 5 to 35 days postpartum
fat and protein contents of beagle milk from (4, 7, 10, 28, 30-33) are considered collec
0 to 40 days, and a subsequent decline in fat tively, a mean ash content of 1.15% can be
content. These trends are at odds with other calculated (n = 57). Assuming the NPN
findings and could reflect sensitivity of the constituents of dog milk to be similar in
spectrophotometric assay procedures to proportion to those of cow's milk (38), NPN
qualitative as well as quantitative changes X 5.34 = 0.29% gives an approximation of
in milk constituents. Canine caseins and the combined weight of NPN constituents
whey proteins certainly differ in amino acid (11). The remaining 0.44 g ( = 1.88 - 1.15
composition (37) such that varying propor - 0.29) represents minor organic and
tions during lactation (6) will affect protein inorganic compounds not included in the
content as measured by dye-binding. various analytical fractions (38) as well as
LACTATION IN THE DOG 809
analytical error. On this basis it would that at least 47% of the isotope lost by dingo
appear that any such error was small. puppies is ingested by the mother; some of
Water and milk intakes. Suckling beagle this is recycled to the young in milk. Isotope
puppies were estimated to consume 142 g recycling was not measured directly in the
water, equivalent to 160 g milk, at 19 days present study although accumulation of
postpartum, and 156 g water, equivalent to D2O in uninjected control puppies indicated
175 g milk, at 26 days postpartum (table 2). that it did occur.
These values are only as accurate as the The milk intakes determined herein are
estimates of body water fraction (F) and much higher than the 81 ±13 ml/day and
water turnover rate (fc)on which the calcu 59 ±13 ml/day reported by Romsos et al.
lations are based. Isotope dilution proce (9) for beagle puppies suckling bitches fed
dures have been reported by Sheng and two semipurified diets. Aside from the
Huggins (23, 25) to overestimate body water methodological problems mentioned above,
content in growing beagles when compared these puppies exhibited abnormally low
to values obtained by direct dessication. growth rates, gaining on average only 13
Dilution of tritiated water indicated body and 11 g/day in the two groups. Normal
per day; i.e., one or two suckling bouts per Weaned puppies have been estimated by
day were relatively unsuccessful. These the Subcommittee on Dog Nutrition, NRC
puppies reportedly consumed 1.7-3.0 g milk (44) to require 22% protein in the dry
per gram body weight gain in weeks 3 and matter of a diet containing 3.5-4.0 kcal ME
4 (7). By contrast puppies in the present per gram dry matter. Dog milk contains
study were calculated to consume 4.4 g milk about 131 kcal ME/100 g or 5.77 kcal ME
per gram gain during week 3 and 4.8 g milk per gram dry matter, a value about 50%
per gram gain during week 4. On this basis above the NRC diet. If dog milk is to meet
it appears that the weight differential pro NRC requirements it should contain 1.5
cedure underestimated milk intake by about x 22 = 33% protein on a dry matter basis.
one-third. The protein content of dog milk was indeed
Peak milk yields of beagle bitches appear found to be 33.4% of dry matter.
to be about 1 kg/day in week 4 postpartum. Puppies ingested 0.33 g protein per gram
Milk yield is undoubtedly influenced by body weight gain at 19 days and 0.36 g
body size (2, 3). Using a weight differential protein per gram body weight gain at 26
procedure, Russe (6) estimated that at 6 days. Payne (45) assumed weight gain in
days postpartum a German shepherd pro puppies to contain 17% protein, whereas
(1960) Composition of beagle dog milk. Am. J. 25. Sheng, H. & Huggins, R. A. (1971) Growth of
Physiol. 199, 731-732. the beagle: changes in chemical composition.
6. Russe, I. (1961) Die Laktation der Hündin. Growth 35, 369-376.
Zentralbl. Veterinaehrmed. 8, 252-281. 26. Ssubotin, Dr. (1866) Ueber den Einfluss der
7. Mundt, H. C., Thomée,A. & Meyer, H. (1981) Nahrung auf die quantitative Zusammensetzung
Zur Energie—und Eiweissversorgung von Saug der Milch. Virchows Arch. A Pathol. Anat. 36,
welpen überdie Muttermilch. Kleintier-Praxis 26, 561-570.
353-360. 27. Tblmatscheff, Dr. (1867) Zur Analyse der Milch.
8. Lönnerdal,B., Keen, C. L., Hurley, L. S. & Fisher, Hoppe-Seylers Med. Chem. Untersuch, l, 272-278.
G. L. (1981) Developmental changes in the 28. Abderhalden, E. (1898) Die Beziehungen der
composition of beagle dog milk. Am. J. Vet. Res. Wachsthumsgeschwindigkeit des Säuglings zur
42, 662-666. Zusammensetzung der Milch beim Kaninchen, bei
9. Romsos, D. R., Palmer, H. J., Muiruri, K. L. & der Katze und beim Hunde. Hoppe-Seylers Z.
Bennink, M. R. (1981) Influence of a low Physiol. Chem. 26, 487-497.
carbohydrate diet on performance of pregnant and 29. Abderhalden, E. (1899) Die Beziehungen der
lactating dogs. J. Nutr. Ill, 678-689. Wachsthumsgeschwindigkeit des Säuglings zur
10. Daggs, R. G. (1931) Studies on lactation. I. Zusammensetzung der Milch beim Hunde, beim
Production of milk in the dog as influenced by dif Schwein, beim Schaf, bei der Ziege und beim
ferent kinds of food proteins. J. Nutr. 4, 443-467. Meerschweinchen. Hoppe-Seylers Z. Physiol. Chem.
11. Oftedal, O. T. (1981) Milk, protein and energy 27, 408-462.