Nutrient Requirements

Milk Intake of Suckling Kittens Remains Relatively Constant from One to

Four Weeks of Age1,2
Wouter H. Hendriks3 and Søren Wamberg*4
Monogastric Research Centre, Institute of Food, Nutrition and Human Health, Massey University, Palmerston
North, New Zealand and *Department of Physiology, Institute of Medical Biology, Odense University,
DK-5000 Odense C., Denmark

ABSTRACT The daily milk intake of 14 domestic short-haired kittens (Felis catus) from five litters was estimated

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during wk 1– 4 postpartum using the isotope dilution technique. Kittens received a single intraperitoneal injection
of tritiated water, and blood samples were obtained from the jugular vein for radioactivity measurements at 2 and
96 h after injection. One kitten in each litter was used as a control to allow calculation of recycling of tritiated water.
The mean (6 SEM) biological half-life of tritiated water in the kittens increased from 2.4 6 0.1 d in wk 1 to 4.9 6 0.2 d
in wk 4 postpartum. Recycling of tritiated water accounted for (mean 6 SEM) 5.9 6 0.8, 12.0 6 0.5, 7.7 6 1.3 and
10.0 6 1.3% of the kittens’ daily water intake during postnatal wk 1– 4, respectively. Daily milk intake of the kittens
during wk 1– 4 postpartum was 47.3 6 0.8, 47.4 6 1.5, 48.7 6 1.6 and 43.7 6 2.0 g, respectively. There was no
effect of gender on milk intake. The daily metabolizable energy requirement of suckling kittens, estimated by
multiple regression analysis, was 356 kJ/kg0.75, whereas the metabolizable energy required per gram of gain was
estimated to be 7.8 kJ/d. The milk intake of suckling kittens remained relatively constant throughout the first 4 wk
of lactation, and during this period, they seemed to have a lower energy requirement for maintenance. J. Nutr.
130: 77– 82, 2000.

KEY WORDS: ● cats ● milk intake ● tritiated water ● water turnover

Knowledge of milk intake by neonatal mammals is of basic
biological interest and is required for the estimation of daily
nutrient intake during this period of an animal’s life. Addi-
tionally, accurate data on milk intake of young animals allow
the milk energy output of the lactating female to be calculated.
Milk intake of suckling young has been measured indirectly
by comparison of growth rates of suckling and formula-fed
young (Buss and Voss 1971), and by weighing the young
several times a day over an extended period before and after
suckling (Mundt et al. 1981, Pettigrew et al. 1985). A more
direct approach for measuring milk intake can be obtained by
the water isotope dilution (WID)5 technique. This method
uses an isotope of hydrogen (such as 2H or 3H) as a tracer to
estimate body water turnover; together with accurate informa-
1
Presented in part at the First European Zoo Nutrition Meeting, 8 –11 January
1999, Rotterdam, The Netherlands [Wamberg, S. & Hendriks, W. H. (1999) Nu-
trient intake of 1– 4 wk old suckling kittens (Felis catus): a model for artificial
rearing of young Felidae. In: Zoo Animal Nutrition No. 1 (Nijboer, J., Hatt, J. M.,
Kaumanns, W., Beynen, A. C. & Ganslosser, U., eds.). Filander Press, Fürth,
Germany].
2
Supported by the Danish Agricultural and Veterinary Research Council,
grant no. 9502267.
3
4
To whom correspondence should be addressed.
Sabbatical visitor at the Heinz Wattie’s Companion Animal Nutrition Re-
search Unit, Institute of Food, Nutrition and Human Health, Massey University,
Palmerston North, New Zealand.
5
Abbreviations used: bwt, body weight; DM, dry matter; MBW, metabolic
body weight; ME, metabolizable energy; MI, milk intake; MWI, milk water intake;
PPO, 2,5-diphenyloxazole; T1/2, biological half-life; THO, tritiated water; TWI, total
water intake; WID, water isotope dilution; WSW, weigh-suckle-weigh.
tion on the composition of milk and body weight gain of the
young, this allows estimation of milk intake (Coward et al.
1982, Fjeld et al. 1988, Pettigrew et al. 1987). A major
advantage of the WID technique over the weigh-suckle-weigh
(WSW) technique is that there is only minimal disruption of
the normal maternal-offspring relationship. Furthermore, the
WSW technique has been shown to underestimate milk con-
sumption by up to 12% in pigs (Pettigrew et al. 1985), 15% in
human infants (Butte et al. 1983), and 30% or more in dogs
(Oftedal 1984) and mice (Baverstock and Elhay 1981).
Recently, Dobenecker et al. (1998) used the WSW method
to estimate milk yield of queens nursing different sized litters
over the first 9 wk of lactation. These authors estimated the
degree of underestimation of the kitten’s milk intake to be
;20 –25%, based on the difference in growth rates during the
measurement period and when suckling normally. Jayawick-
rama et al. (1998) used the WSW technique and reported milk
intake of 20 –32 and 54 –71 g/d by kittens suckling on queens
fed diets containing 10 and 20% fat, respectively. To our
knowledge, direct measurements of milk intake in kittens
during the suckling period using the isotope dilution technique
are not available.
Given the known problems associated with the WSW
method, we decided to use the isotope dilution technique to
increase the accuracy of estimates of milk and nutrient intake
in suckling kittens. Milk intake of kittens during the first 4 wk
of life was studied using tritiated water (THO) as a marker.
Furthermore, at the end of the main study, the accuracy of the

0022-3166/00 $3.00 © 2000 American Society for Nutritional Sciences.

Manuscript received 21 May 1999. Initial review completed 3 July 1999. Revision accepted 17 August 1999.

77
78 HENDRIKS AND WAMBERG

WID technique in estimating milk intake was assessed by tube TABLE 1

feeding six kittens over a period of 48 h.
Nutrient composition of the diet and milk fed to the queens
and the reconstituted kitten milk replacer
MATERIALS AND METHODS
Queen
The studies reported here complied with the Massey University Kitten milk
Code of Ethical Conduct for the use of Live Animals for Teaching Ingredient Diet1 Milk2 replacer3
and Research (Anonymous 1992).
Animals, housing and diets. Five lactating domestic short- g/kg as is
haired queens (Felis catus) and their offspring were maintained in
standard metabolism cages (described in Hendriks et al. 1999) nor- Dry matter 18.3 13.7 8.2
mally used for the rearing of kittens in a closed colony (Heinz-
Wattie’s Companion Animal Nutrition Research Unit, Massey Uni- g/kg dry matter
versity, Palmerston North, New Zealand). All queens had been
vaccinated against feline rhinotracheitis, calicivirus and panleukope- Crude protein 493 347 409
nia using a modified live vaccine (Felocell CVR, Norden Laborato- Lipids 283 234 186

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ries, München, Germany). Feline leukemia and feline immunodefi- Ash 93 52 43
ciency virus have not been detected in the colony since its Taurine 3.1 0 0.3
establishment in 1976. One queen gave birth to three kittens and four Methionine 12.3 6.5 12.3
queens gave birth to four kittens. The range in body weights of the 19 Aspartic acid 41.4 19.1 31.7
kittens (10 males and 9 females), measured within 24 h after birth, Threonine 20.9 11.3 19.8
was 100 –140 g (mean 6 SEM, 120 6 2 g). On postnatal day 3, the Serine 21.0 13.6 23.6
Glutamic acid 64.7 48.6 94.0
kittens in each litter were weighed accurately and a sterile microchip
Glycine 43.2 4.9 7.7
(Life Chip, Animal ID Electronic Systems, Kiama, NSW, Australia) Alanine 32.5 7.8 12.4
was inserted subcutaneously for individual identification before sub- Valine 25.6 16.0 29.1
sequent experimental procedures such as weighing, injection of triti- Isoleucine 17.9 12.7 22.0
ated water and blood sampling. The queens were given free access to Leucine 40.9 24.3 41.7
a homogenized mixture of moist canned cat foods and lactose-free Tyrosine 16.4 11.6 21.5
milk throughout the study. Table 1 shows the analyzed composition Phenylalanine 21.7 12.1 22.0
of the mixture of canned cat foods and the lactose-free milk. Fresh Histidine 15.9 7.0 12.6
drinking water was available at all times. The body weights of the Lysine 30.4 19.3 34.6
queens and kittens were recorded at regularly preset intervals Arginine 30.1 8.4 14.6
throughout the study.
Experimental procedures. The queens and their offspring were 1 Mixture of commercial canned cat foods (Chef, Heinz Wattie’s,
considered healthy on the basis of clinical examination and hema- Auckland, New Zealand).
tologic status before the start of the study. In the morning of postnatal 2 Pets Own lactose free milk, Appetite Foods (Lysterfield, Victoria,
days 3, 10, 17 and 24, each kitten was weighed and given an 3156, Australia).
accurately weighed amount (;2 mL/kg body weight) of sterile iso- 3 Made up from a mixture of 33.5 g goat’s milk powder (Karicare
tonic (9 g/L) saline containing 25 mCi (925 MBq)/L of THO (Life Goat Follow-On, Nutricia, Auckland, New Zealand) and 16.5 g sodium
Sciences Technologies, Auckland, N.Z.) by intraperitoneal injection. caseinate (Alanate 180, New Zealand Dairy Board, Wellington, New
In each litter, one blank kitten served as a control for the calculation Zealand) in 500 mL demineralized water.
of recirculation of THO, due to the uptake of kittens’ urinary and
fecal water by the queen (Baverstock and Green 1975); that kitten,
therefore, received only an intraperitoneal injection of sterile isotonic Chemical analysis. Dry matter (DM) of plasma samples was
saline. After injection, the kittens were physically separated from the determined in duplicate by freeze drying and subsequent oven drying
queen for 2 h to allow full equilibration of the injected isotope and to at 105°C. Dry matter of the diets and the milk replacer was deter-
prevent any water exchange between the queen and the kittens. After mined in duplicate by desiccation at 105°C. Ash was determined by
the 2-h equilibration period, a blood sample from the jugular vein was heating samples at 550°C for 16 h. Total nitrogen was determined
taken from each kitten using a 1-mL syringe mounted with a 27-gauge using the Kjeldahl method with crude protein calculated by multi-
needle; the kittens and queen were then returned to the metabolism plying total nitrogen by 6.25. Lipid was determined by petroleum
cage. A further blood sample of each kitten was obtained 4 d after ether extraction of freeze-dried samples (AOAC 1980). Amino acids
injection. Handling of the kittens was kept to a minimum, and efforts were determined as described by Hendriks et al. (1996). All measure-
were made to reduce the disturbance of the mother-young relation- ments were performed in duplicate and the chemicals used were
ship. analytical grade.
Measurement of blood plasma THO. Each blood sample (0.2– Total body water and biological half-life of body water turnover.
0.4 mL) of a kitten was immediately transferred into a 0.5-mL The body water content of the kittens at various ages, which is
heparinized Eppendorf tube. The blood was then used to fill 3–5 required in the calculation of water intake, was estimated using the
standard (75-mL) hematocrit capillary tubes and centrifuged at following regression line [values are: estimate (6SEM)]:
14,800 3 g for 15 min using a Heraeus hemofuge (Heraeus Sepatech
GmbH., Osterode, Germany). The hematocrit values were recorded; Total body water ~%! 5 79.9 ~60.4! 2 0.22
then the capillary tubes were cut, and the plasma was used to fill two ~60.03! z age ~d! ~r 5 0.81, n 5 28) (1)
50-mL calibrated micropipettes (Vitrex, Hounisen, Risskov, Den-
mark). Each micropipette was transferred directly into a scintillation The regression line was obtained by least-squares regression of the
vial containing 8 mL of scintillation fluid [67 parts toluene and 33 water content of newly born to 6-wk-old kittens published by Thomas
parts triton X-100 containing 0.4% 2,3-diphenyloxazole (PPO)], and (1911), Widdowson (1950) and Stratmann (1988). The biological
the radioactivity was counted using a Wallac 1414 liquid scintillation half-life (T1/2) of body water in the kittens was calculated from rates
spectrometer (Wallac OY, Turku, Finland). Any remaining plasma of elimination from the body water pool of the injected THO over
samples of the kittens were pooled per period (3–7, 10 –14, 17–21 and the 4-d observation period. The tritium radioactivity in plasma water
24 –28 d postpartum) for dry matter analysis. Standards were made up at 2 and 96 h after injection of THO and the equations published by
by dilution (1:1000) with distilled water of the THO solution used for Coward et al. (1982) were used to calculate water intake in THO-
injection, and corrections were made for the quenching effect of the injected kittens. Recycling of THO between the queen and her
plasma samples. The average counting efficiency of tritium was 42%. kittens during wk 1 was accounted for by subtraction of the plasma
 MILK INTAKE REMAINS CONSTANT IN GROWING KITTENS
TABLE 2
Weekly body weight, metabolizable energy (ME) intake and milk yield of the queens and body weight and weight gain of the
kittens during the study period1
1 2
Queens, n 5 5
Body weight,2 kg 3.31 6 0.13
ME intake, kJ/(kg z d) 392 6 23
Milk output3, %/kg 5.1 6 0.3
Milk output, %/kg0.75 7.0 6 0.4
Kittens, n 5 19
Body weight,1 g 214 6 3
Body weight gain, g/d 15.0 6 0.4
1 Values are mean 6 SEM.
2 Weight on the last day of the week.
3 Milk yield in percentage of queen’s body weight.
water counts of the blank kitten at 96 h from the corresponding
individual values of the THO-injected kittens. Recycling of THO
during wk 2, 3 and 4 was calculated similarly, with the recycling of
THO determined from the difference between calculated and mea-
sured plasma counts of the blank kitten at 96 h. The calculated
plasma THO radioactivity was determined using the plasma counts at
2 h and the average fractional rate of water turnover of the THO-
injected kittens during the previous period.
Calculation of milk intake. The daily milk intake was calculated
from the water intake data using the equations published by King et
al. (1993). Because the changes in the composition of the major
constituents in queen’s milk appear to be rather small within the first
4 wk of lactation (Adkins et al. 1997, Dobenecker et al. 1998), and
because such changes cause only very small errors in the calculated
value for daily milk intake (Pettigrew et al. 1987), the following
composition of queen’s milk (compiled from literature values) was
used for calculations in this study: 79.4% water, 8.2% crude protein,
5.5% fat and 5.5% lactose. The digestibility of milk protein, fat and
lactose was assumed to be close to 100% (Kienzle and Kamphues
1991). Full oxidation of one gram of protein, fat and lactose in vivo
was taken to yield 0.41, 1.07 and 0.60 mL of water, respectively
(Brody 1945). Potential metabolic water deposited as protein and fat
(King et al. 1993) was calculated using the body dry matter gain
(DMgain) of each kitten over the measurement period and values of
62 and 22 g/100 g dry matter for body protein and fat content of the
kittens, respectively (Stratmann 1988). Body dry matter gain over the
measurement period was determined using the body weight gain over
each experimental period (4 d) and the average body water content
of kittens at the median of the observation period. The latter value
was calculated using the regression equation (Eq. 1) given above. The
following presents the final equation whereby the daily milk intake
(MI) of the kittens was calculated from total water intake (TWI):
MI ~ g/d! 5 1.088 z TWI ~ g/d! 1 0.520 z DM gain (g/d) (2)
Statistical analysis. The body weight data of the kittens were
subjected to repeated-measures ANOVA with litter and gender as
variables and days postpartum as the repeated measure (Cody and
Smith 1987). The milk intake data were subjected to ANOVA
(split-plot) using the General Linear Model with queen, gender, sex
and kitten as fixed variables and kitten within queen and sex as the
error term to account for the repeated measurements of milk intake
(Cody and Smith 1987). The data on the biological half-life of body
water were subjected to a least-squares linear regression analysis with
body weight as the independent variate and the biological half-life as
the dependant variate. A no-intercept multiple regression analysis
was performed on the milk intake data with milk intake (g/d) as the
dependent variable and metabolic body weight (kg0.75) and body
weight gain (g/d) as the independent variates. All statistical analyses
were performed using the SAS statistical package (SAS version 6.12,
79
Week
3 4
3.05 6 0.12 2.91 6 0.10 2.60 6 0.13
459 6 24 513 6 59 477 6 70
5.5 6 0.3 6.1 6 0.4 6.0 6 0.4
7.4 6 0.3 8.0 6 0.5 7.6 6 0.5
284 6 5 347 6 7 382 6 11
9.5 6 0.5 8.1 6 0.7 3.4 6 0.8
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SAS Institute, Cary, NC), and effects were considered significant at
P , 0.05. Values in the text are means 6 SEM.
Validation of the THO method for measuring milk intake in
growing kittens. At the end of the 4-wk study, six kittens (one male,
one female from each of three litters) were given an intraperitoneal
injection of THO in isotonic saline as described above and separated
from the queen for 2 d in an insulated nest box. After a 2-h equilibration
period, the kittens were fed a freshly prepared milk replacer by stomach
tube (Baxter Feeding Tube no. K31, Baxter Healthcare, Deerfield, IL) for
48 h to test the accuracy of the water isotope dilution technique in
determining milk intake in growing kittens. The kittens were fed a
known amount of warm (640°C) milk replacer three times a day at an
energy level allowing minimal body weight gain to minimize the risk of
diarrhea due to overfeeding. The exact amount of milk replacer was
determined by accurately weighing the syringe 1 stomach tube imme-
diately before and after each feeding. Blood sampling and THO analysis
were as described above. Daily milk intake was calculated using the
composition of the milk replacer shown in Table 1 and compared with
the amount given by stomach tube.
RESULTS
The kittens and queens remained healthy throughout the
4-wk experimental period. The kittens were not observed to
leave their nest box and were never found in the area where
the food for the queen was presented during the study. The
kittens gained weight throughout the study although body
weight gain decreased as the study progressed (Table 2). There
was a significant (P , 0.001) effect of time and no significant
(P . 0.05) effect of litter and gender on the body weight of the
kittens as determined by repeated-measures ANOVA. There
was a significant (P , 0.001) interaction between litter and
time on the body weights of the kittens. The queens lost
weight throughout the study, particularly during wk 3 and 4
postpartum. The average daily intake of ME of the queens
increased during the first 3 wk of the study from 392 kJ/kg body
weight during wk 1 to 513 kJ/kg body weight during wk 3.
During wk 4, the daily ME intake of the queens decreased to
477 kJ/kg body weight.
The hematocrits of the kittens decreased throughout the 4-wk
study period, with the largest decline occurring in the first 2 wk
(Fig. 1). At the end of the study, the hematocrit of the kittens
was 0.19 6 0.01. The dry matter content of the kittens’ plasma,
as measured using the pooled samples, remained constant
throughout the 4-wk study (6.8 6 0.1%). Recirculation of THO
as measured at 96 h after injection during wk 1–4 accounted for
80 HENDRIKS AND WAMBERG
FIGURE 1 Hematocrits of the kittens throughout the 4-wk exper-
imental period. Values are means 6 SD, n 5 19.
5.9 6 0.8, 12.0 6 0.5, 7.7 6 1.3 and 10.0 6 1.3% of the daily
water intake, respectively. Figure 2 shows the linear regression
line relating the biological half-life of body water to the body
weight (bwt) of the kittens during the 4-wk experimental period.
The following significant regression line was obtained [values are:
estimate (6SEM)]:
T 1/2 body water (d) 5 0.90 (60.33) 1 9.52
~61.08! z bwt ~kg! ~r 5 0.76, n 5 56) (3)
The slope and the intercept of the regression line were signif-
icant at P , 0.01. The biological half-life of body water
increased during wk 1– 4 and was 2.4 6 0.1, 3.4 6 0.1, 4.0
6 0.1 and 4.9 6 0.2 d, respectively.
Milk intake of the kittens remained relatively constant
throughout the study (Fig. 3) although there were significant
(P , 0.05) effects of litter, time and the interaction between
litter and time on milk intake. There was no effect of gender
on milk intake. Daily milk intake of the kittens (n 5 14)
during wk 1– 4 were 47.3 6 0.8, 47.4 6 1.5, 48.7 6 1.6 and
43.7 6 2.0 g, respectively. Multiple regression of the milk
intake data on metabolic body weight (MBW) and body
weight gain (gain) yielded the following significant (P
, 0.001) equation [values are: estimates (6SEM)]:
MI ~ g/d! 5 77.9 ~ 6 2.4! z MBW ~kg 0.75)
1 1.7 ~ 6 0.1! z gain ~ g/d! ~r 5 0.73, n 5 56) (4)
Both coefficients of the multiple regression line were signifi-
cant (P , 0.001).
FIGURE 2 Biological half-life of tritiated water turnover in suckling
kittens (n 5 14) measured over 4-d periods during postnatal wk 1– 4
and plotted as a function of body weight (r 5 0.76, P , 0.05).
FIGURE 3 Milk intake of individual kittens (n 5 14) during the four
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experimental periods. Horizontal bars indicate mean values. Means (6
SEM) for wk 1– 4 were 47.3 6 0.8, 47.4 6 1.5, 48.7 6 1.6 and 43.7 6 2.0,
respectively.
Four of the six kittens used in the validation study (wk 5)
gained weight (mean: 5.5 g/d), whereas two kittens lost body
weight (mean: 21 g/d). Diarrhea was not observed during the
48-h period when the kittens were fed the milk replacer. The
daily intake of milk replacer, estimated using the WID tech-
nique, ranged from 0.3% underestimation to 7.2% overestima-
tion with an average overestimation of daily milk intake of 2.4
6 1.2%.
DISCUSSION
All of the queens and kittens in this study remained
healthy, and the average weight gain of the kittens, recorded
during the first 3 wk postpartum, was comparable to the data
of Loveridge (1987) for a total of 100 kittens. In wk 4, the
average weight gain of the kittens was lower than the values
reported by Loveridge (1987), which could be attributed to the
failure of two kittens to gain body weight. At the end of the
study, however, the average body weight of the kittens was
higher than the average body weights (373 6 5 g, n 5 128) of
kittens of a similar age recorded in the colony. The failure of
two kittens to gain body weight in the final week of the study
was due to a low milk intake (75 and 70 g per kg MBW).
There was no effect of litter or gender on the body weight of
the kittens in this study as analyzed by repeated-measures
ANOVA. Loveridge (1987) also found no effect of gender on
the body weight of kittens until 6 wk of age; the male kittens
were significantly heavier after that time. The body weight of
the queens in this study decreased throughout lactation and at
the end of wk 4, the average body weight of the queens was
reduced to 79% of the value recorded at the start of lactation.
The mean daily energy intake of the queens in this study was
similar to that reported by Loveridge (1986), except for wk 4
when a decrease in the intake of ME was recorded. At the
same time, the queens experienced heavy weight loss, indicat-
ing that body reserves were mobilized to provide energy and
nutrients to maintain milk production. In this situation, the
reduced intake of ME by the queens is likely to have caused a
lower milk production, resulting in a lower weight gain of the
kittens in the last week of the study.
In the kittens the decreasing hematocrits observed through-
out the study (Fig. 1) document the development of “suckling
anemia,” a phenomenon that has been observed in rapidly
growing suckling young of other species (Garcia 1957, Wid-
dowson 1965). By measuring iron levels in the body of suck-
ling kittens, McCance and Widdowson (1951) showed that
 MILK INTAKE REMAINS CONSTANT IN GROWING KITTENS
blood iron levels decreased from birth to 3 wk of age. Garcia
(1957) showed that the rapid body growth of suckling rats
leads to a marked decrease in the hematocrit value and the
hemoglobin concentration of whole blood during early post-
natal life. This occurred in spite of a marked increase in total
red cell volume and whole-body hemoglobin, indicating a
continual high rate of erythropoiesis, exceeded by higher rates
of body fluid accretion and body mass gain (Garcia 1957). The
repeated blood sampling in this study also affected the hema-
tocrits, but the effect is believed to be of minor importance
because the amount of blood taken (0.2– 0.4 mL) represented
only a small fraction of the kittens’ total blood volume.
In applying the WID technique to studies of water turnover
in the mammalian body, a number of basic assumptions have
to be fulfilled, including water isotope equilibration, size of the
body water pool, sources and rates of water exchange and loss
of isotopes. The assumptions, prerequisites and the potential
errors inherent in using this technique for determinations of
water fluxes in animals and humans and its implications for
reliable estimates of the milk intake in various species have
been reviewed (Coward et al. 1982, Fjeld et al. 1988, Nagy and
Costa 1980). In this study, a 2-h period for THO equilibration
was used because this seems to be a commonly accepted
equilibration time for water isotopes in small mammals weigh-
ing ,10 –20 kg (Fjeld et al. 1988, Macfarlane et al. 1969,
Oftedal 1984, Pettigrew et al. 1985, Pluske et al. 1998). At the
end of the study, an equilibration time curve was obtained
from the use of six kittens injected with THO, and it was
shown that full equilibration of THO in kitten’s body water
was attained in ,1 h (Hendriks, unpublished observations). In
this study, the recirculation of THO due to the uptake of urine
and feces by the queen was measured in unlabeled kittens
during wk 1; these values, therefore, can be expected to be
accurate. To allow repeated measurements on the same litter
and take into account recirculation of THO, a previously
THO-injected kitten was used as a “blank” during wk 2 to 4.
In these cases, the calculations of recirculated THO were
corrected for residual THO radioactivity in plasma water from
the previous measurement period, and the calculated values
for wk 2– 4, therefore, are theoretically slight overestimates.
THO recirculation values found for wk 1– 4 were 5.9 6 0.8,
12.0 6 0.5, 7.7 6 1.3 and 10.0 6 1.3%, respectively.
The biological half-life of body water in the kittens in-
creased from 2 d in wk 1 to 4 d in wk 4 (Fig. 2); the latter value
is similar to that observed in 3-wk-old suckling puppies (4.2–
4.6 d) (Oftedal 1984). In suckling mink kits, weighing 20 –140
g, the biological half-life increased from ,1 d to ;2 d (Wam-
berg and Tauson 1998), whereas in 1- to 3-wk-old blue fox
cubs (Alopex lagopus), the T1/2 value of body water ranges from
1.7 to 2.2 d (Wamberg, unpublished). In suckling rat pups,
T1/2 is ;1.5 d (Coward et al. 1982); in human infants weigh-
ing 7.8 kg, Fjeld et al. (1988) reported a mean value of T1/2 of
2.7 d. These values underscore the rapid turnover rate of body
water in suckling young animals.
The isotope dilution technique has been used extensively
to measure milk intake in a number of mammalian species,
including humans (Butte et al. 1983), baboons (Buss and Voss
1971), sheep (Macfarlane et al. 1969), rats (Coward et al.
1982), dogs (Oftedal 1984), pigs (Pluske et al. 1998) and mink
(Wamberg and Tauson 1998). In this study, we have applied
the WID technique for the first time to study milk intake in
suckling kittens and determine the accuracy of this technique
to measure milk intake in kittens. The measurement of milk
intake by the WID technique in the tube-fed kittens showed
a high degree of accuracy. The mean difference between the
predicted and actual milk intake of the six measurements was
81
2.4%. This value is similar to the recovery values reported in
infants of 2.0% (Fjeld et al. 1988), calves 2.5% (Holleman et
al. 1975), lambs 3.3% (Macfarlane et al. 1969), and pigs 2.9%
(Pettigrew et al. 1987). These results indicate that the results
presented on the milk intake of kittens in this study were
accurate.
The average daily milk intake of the kittens remained
surprisingly constant throughout the first 4 wk postpartum
(Fig. 3). However, there was a significant effect of time on
milk intake and a significant interaction between milk intake
and litter, indicating that milk production patterns of the five
queens were different over time. The body weight gain of the
kittens also showed a significant effect of time; the kittens
were found to grow at a different rate as indicated by the
significant interaction of time and litter. Jayawickrama et al.
(1998) found a significant difference between the milk intake
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of kittens for wk 1 and 4 and statistically similar milk intakes
among other weeks. Dobenecker et al. (1998) also used the
WSW technique to measure milk intake of kittens during
three periods (wk 1, 2– 4, 5–9) and found a higher milk yield
of queens during the wk 2– 4 period compared with wk 1 and
a lower milk yield during the wk 5–9 period. The results in this
study are in contrast to the increasing milk intake with in-
creasing age observed in the suckling young of baboons (Buss
and Voss 1971), rats (Coward et al. 1982), dogs (Oftedal
1984) and mink (Wamberg and Tauson 1998). In pigs, Pluske
et al. (1998) found a significant decrease in milk intake with
increasing age. Many factors may influence milk production
throughout lactation in mammals, including factors such as
stage of lactation, the number of suckling animals, suckling
intensity, temperature, season, dietary protein and energy in-
take (Dobenecker et al. 1998, Jayawickrama et al. 1998, King
et al. 1993, Pluske et al. 1998).
Milk intake and the growth of the suckling offspring are
highly correlated because milk is the sole source of nutrients
for the young animal. The milk ingested by the kittens in this
study would have been used first for the maintenance of body
mass; any intake of milk in excess of maintenance would have
been used for growth of body tissue. The average body weight
of the kittens in this study increased with age, whereas the
growth rate decreased with age (Fig. 2), indicating that the
amount of milk available for body gain decreased throughout
the 4-wk lactation period. The multiple regression equation
obtained in this study (Eq. 4) shows that 77.9 g of milk were
required per unit metabolic body weight per day for mainte-
nance and 1.7 g were required per gram of body growth.
Assuming that milk energy contains 4.6 kJ ME/g, this equates
to suckling kittens requiring 356 kJ/(kg0.75zd) for maintenance.
This value is close to the basal metabolic rate of kittens [288
kJ/(kg0.75zd)] as measured by oxygen uptake of 1- to 6-wk-old
kittens (Hill 1959). Similar values [300 and 334 kJ/(kg0.75zd)]
for maintenance have been found in suckling dogs (Crighton
and Pownall 1974, Mundt et al. 1981). This value, however, is
considerably lower than values normally found for pigs of 458
kJ/(kg0.75zd) (Lawrence and Fowler 1997) and adult cats (NRC
1986) and dogs (NRC 1985) of 438 and 552 kJ/(kg0.75zd),
respectively. Kittens and dog puppies are born with a dense
hair coat and exhibit very low physical activity because they
sleep most of the day. They huddle together and are kept warm
by the queen or bitch most of the time. All of these factors will
reduce the energy requirement for maintenance to a level
closer to the basal metabolic rate of the animal. The efficiency
of ME deposition can be calculated using the data in this
study. Per gram of body growth, 1.7 g of milk or 7.8 kJ ME is
required per day. Assuming that the body weight gain con-
sisted of 76.6% water, 14.4% protein and 5.8% fat (Stratmann
82 HENDRIKS AND WAMBERG
1988), the energy stored per unit of growth was 5.5 kJ/d. The
efficiency (kg-value) with which metabolizable energy was
converted to net energy in the kittens in this study was 0.71.
This is similar to other animals such as pigs, in which kg-values
of 0.65 and 0.69 have been reported (Lawrence and Fowler
1997, Van der Hel and Verstegen 1987). It has been noted by
several authors that in doubling its body weight in ,8 d, the
cat is one of the fastest growing mammals (Bernhart 1961,
Thomas 1911, Widdowson 1965). This study indicates that
the high growth rate of cats may be due mainly to the lower
energy requirement for maintenance, ;360 kJ/(kg0.75 z d),
whereas the ME requirement per unit of body growth is similar
to that of other mammals.
The average daily milk production of the queens in this
study (as a percentage of the queens’ body weight) ranged from
5.1 6 0.3% during wk 1 to 6.1 6 0.4 and 6.1 6 0.4% during
wk 3 and 4, respectively. Dobenecker et al. (1998) measured
milk production of queens using the WSW technique and
reported corrected estimates of milk production of queens
nursing litters of 3– 4 kittens during wk 1 of 4.0% and during
wk 2– 4 of 5.7%. However, the expression of the daily milk
yield as a percentage of body mass may be misleading because
queens may lose a considerable amount of body mass during
lactation. Estimates made by Dobenecker et al. (1998) for wk
1 of lactation are lower than the estimates obtained in this
study, and their data show a distinct relationship in the milk
yield and lactation stage, which is likely to be caused by the
WSW technique used to measure milk intake. The differences
between these two techniques in estimating milk intake have
been attributed to stress resulting from repeated interference
and frequent handling of the young, to lack of suckling stim-
ulus, small errors in frequent weighings, loss of excreta and to
the effects of water recycling (Baverstock and Elhay 1981,
Coward et al. 1982, Oftedal, 1984, Pettigrew et al. 1987).
Similarly, in recent studies of the milk intake of suckling
kittens using the WSW technique, Dobenecker et al. (1998)
and Jayawickrama et al. (1998) found lower growth rates of the
kittens during the measurement period than those of the same
kittens in periods in which they were not separated from the
queens. These lower growth rates are likely to have resulted
from stress of the queens and kittens, lower milk intake of the
kittens, lower milk output by the queen, water recirculation or
a combination of these factors.
ACKNOWLEDGMENT
The authors thank H. Nicol for the skilled care of the cats
throughout this study.
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