Waltham International Symposium:
Pet Nutrition Coming of Age
Influence of Calcium and Phosphorus Intake on the Apparent Digestibility
of These Minerals in Growing Dogs1,2
Britta Dobenecker3
Institute for Physiology, Physiological Chemistry and Animal Nutrition, Ludwig-Maximilians Universität,
Munich, Germany
EXPANDED ABSTRACT
KEY WORDS: ● symposium ● dogs ● developmental skeleton disease
Skeletal diseases are often diagnosed in growing dogs, most
commonly in large and giant breeds. The connection between
mistakes in feeding regime and disturbances of bone develop-
ment has been illustrated by the negative effects of a lack in
calcium (Ca4)4 intake (1) and the consequences of an over-
supply of energy and/or Ca for skeletal health (1– 4). The
inability of puppies to decrease the digestibility of Ca to
compensate for a threefold oversupply has already been shown
by Hazewinkel and coworkers (3). In the literature there is
only limited information about the digestibility of Ca and
phosphorus (P) in puppies with an insufficient intake of those
minerals, especially regarding the ability to make optimum
feeding recommendations after a period of malnutrition in
growing dogs.
MATERIALS AND METHODS
To determine and quantify the ability of growing dogs to alter the
apparent digestibility (aD) of Ca and P at various degrees of over- and
undersupply at different ages during growth, a feeding trial with 20
healthy Beagles from three litters (10 m/10 f) was carried out from
weaning until 6 mo of age. Four groups were formed to carry out three
trials of approximately 6-wk duration each (42 to 49 d). The dogs of
each group were housed together in a large kennel with access to an
outdoor area. A diet based on tripe, rice and cellulose and supple-
mented individually with minerals and vitamins was fed (Table 1).
1
Presented as part of the Waltham International Symposium: Pet Nutrition
Coming of Age held in Vancouver, Canada, August 6 –7, 2001. This symposium
and the publication of symposium proceedings were sponsored by the Waltham
Centre for Pet Nutrition. Guest editors for this supplement were James G. Morris,
University of California, Davis, Ivan H. Burger, consultant to Mars UK Limited, Carl
L. Keen, University of California, Davis, and D’Ann Finley, University of California,
Davis.
2
Supported by Gesellschaft zur Förderung Kynologischer Forschung, Ger-
many.
3
To whom correspondence should be addressed.
E-mail: Dobenecker@lrz.uni-muenchen.de.
4
Abbreviations used: aD, apparent digestibility; Ca, calcium; DM, dry matter;
f, female; g, gram; kg, kilogram; m, male; n, number of animals; P, phosphorus;
tD, true digestibility.
0022-3166/02 $3.00 © 2002 American Society for Nutritional Sciences. J. Nutr. 132: 1665S–1667S, 2002.
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● minerals ● digestibility
The amount of the basic ration was given based on the energy
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requirement of the dogs (5) and the energy content in the diet (6).
The intake of Ca, P and the other minerals and vitamins was
determined and balanced individually and exactly with the help of a
special supplement for all groups: every single dog of each group got
an especially mixed mineral and vitamin supplement, which group
was given according to actual body weight and age. Depending on the
sources of Ca and P in the supplements were CaCO3, Ca-phosphate
and Na-phosphate. All but the nutrients in question were supplied to
meet or exceed the requirements per kg body weight (5).
The course of trials was as follows (Table 1). The Ca intake was
adjusted to 15, 50, 100 (control), 150 and 300% of the requirement
(the Ca and P content in the control diet at the beginning of
Trial 1 was approximately 11 g Ca and 8 g P/kg DM; and at the end
of Trial 3 approximately 10 g Ca and 7 g P/kg DM). The P intake met
the requirements, that is, the Ca/P ratio was altered accordingly when
the Ca intake was adjusted. After Trial 1 and without a break in
between, the feeding of groups A to C was changed to Trial 2 diets,
from oversupply to undersupply and vice versa. In both trials the five
dogs of the control group were fed according to the requirements
(100%) (i.e., all nutrients met the requirements). After Trial 2 all
dogs were changed to a 100% diet for another 6 wk (Trial 3). At the
end of each feeding period a balance period of 7 d was carried out;
thus there was a washout period of a minimum of 5 wk before the
balance period. During the balance period the dogs were housed
separately in cages that allowed sampling of feces and urine. The feces
were collected quantitatively, then weighed, lyophilized and ground
for further analysis. The Ca content was measured with flame emis-
sion photometry (7) and the P content by photometry after acid
hydrolysis (8). Body weight was determined every day before feeding.
Data are expressed as means ⫾ SD. Means were compared by
Tukey’s test to determine significant differences between the groups
(P ⬍ 0.05 was considered a significant difference). The study was
approved by the Regierung of Oberbayern, which is the proper
authority according to German laws on animal welfare (Tierschutzge-
setz).
RESULTS AND DISCUSSION
As intended, no impairment of bone health was evident: no
clinical symptoms of disturbances of skeletal development
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TABLE 1 group (300%) relative to the control group. In the case of a

high Ca intake there is little difference between aD and true
Composition and average nutrient content (% dry matter) digestibility (tD) because endogenous losses can be assumed to
of the basal diet fed to 20 growing Beagles be relatively small when compared to the levels of intake and
fecal excretion. In addition, endogenous fecal losses have been
Component Amount (%)
repeatedly estimated by regression analysis and they appear to
Basal ration be fairly constant in dogs (9). If the Ca intake is low, the aD
Tripe 60 is considerably lower than the tD. Therefore it is most unlikely
Cooked rice 39 that a significantly higher aD in a group with reduced Ca
Cellulose 1 intake is an effect of endogenous losses. In contrast, the
Supplement
Vitamin-mineral mixture Individual supplementation
differences in the tD are likely to be more distinct than those
Average nutrient content, % dm in the aD. Trial 3 showed no significant differences in the aD
Dry matter 31.2 ⫾ 3.7 of Ca. At this age the digestibility seemed to be unaffected by
Crude protein 33.7 ⫾ 5.6 the preceding feeding, which makes the influence of carryover
Crude ash 2.3 ⫾ 0.6 effects less likely.
Crude fat 28.8 ⫾ 8.9
Crude fiber 5.1 ⫾ 1.9
In contrast to the influence of the diets on the aD of Ca, the
N-free extracts 31.0 ⫾ 11.9 aD of P was influenced significantly in the youngest dogs (Trial
1). When the Ca intake or the Ca/P ratio did not meet the
requirements as in groups A to C, a difference in the values
was measured. The aD of P was significantly lower in the case
were noticeable in any of the Beagle puppies. The weight of a Ca oversupply in Trial 1, as Jenkins and Phillips (10) have

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curve was not influenced systematically by the feeding regime.
In contrast to the aD of Ca, which was highly variable and
showed only a tendency to decrease, the aD of P decreased
significantly with the age in the control group (100%), as
expected (9); so the highest values were measured in the
youngest puppies. At the age of 6.5 to 13.5 wk of age (Trial 1)
no statistically significant differences in the aD of Ca were
seen among the groups (Table 2). This shows that during this
age the puppies are not able to adjust the aD according to the
intake, as has already been shown by Hazewinkel and cowork-
ers (3) in the case of an oversupply with Ca. The first time a
difference between the groups could be measured was in older
pups, that is, in Trial 2 (13.5–19.5 wk of age). Here, in the
15% group (group B), a significant higher digestibility of Ca
compared to the 300% group (group A) was measured. Neither
the control group (100%) nor group C, which was fed with a
ration to meet 50% of the Ca requirements, was significantly
different from the other two groups. Consequently, no down-
regulation of the aD of Ca was detectable in the oversupplied
already described. A distinctly higher mean value for the aD of
P was detectable in puppies on a Ca undersupply (15%) and a
Ca/P ratio far below 1:1 compared to that of the other groups.
A potential mechanism for this observation might be a possi-
ble elevation of the blood level of PTH and, as a consequence,
a stimulation of the kidneys to form active vitamin D, which
leads to a higher absorption of both elements from the gastro-
intestinal tract. Trial 3 showed no significant differences in the
aD of P.
The results demonstrate that dogs younger than 4 to 5 mo
are not able to adjust the digestibility of Ca in relation to the
Ca intake at 100% of requirements. Not only the inability to
downregulate the aD but also the inability to increase the
uptake in the case of an insufficient intake in young growing
dogs makes it necessary to adjust precisely the daily supply of
Ca to the requirements. In contrast, an ability to alter the
apparent digestibility of P was already observed in very young
dogs.

TABLE 2
Apparent digestibility (aD, mean ⫾ SD, %) of calcium and phosphorus in 20 Beagles (from weaning to 6 mo of age)
in relation to age and daily intake of these elements

Trial Feeding regime

(age of dogs, wk) Group n (% ca) Ca/P aD Ca aD P

1 (6.5–13.5) Control 7 1001 1.5/1 65.6 ⫾ 8.4 79.5*a ⫾ 3.0

A 5 15 0.2/1 46.2 ⫾ 32.9 93.4b ⫾ 1.1
B 5 300 4.8/1 54.1 ⫾ 10.5 73.1c ⫾ 2.7
C 3 150 2.4/1 47.7 ⫾ 3.5 66.9d ⫾ 2.0
2 (13.5–19.5) Control 6 100 1.6/1 56.0ab ⫾ 15.9 65.5*a ⫾ 11.7
A 5 300 5.0/1 52.0b ⫾ 9.3 62.5a ⫾ 10.3
B 5 15 0.2/1 73.8a ⫾ 9.3 92.0b ⫾ 0.4
C 3 50 0.9/1 57.3ab ⫾ 0.8 72.2a ⫾ 1.3
3 (19.5–25.5) Control 6 100 1.6/1 43.3 ⫾ 20.1 62.2* ⫾ 9.0
A 5 100 1.6/1 55.7 ⫾ 12.2 71.4 ⫾ 8.6
B 5 100 1.6/1 54.4 ⫾ 9.4 69.8 ⫾ 6.4
C 3 100 1.6/1 45.0 ⫾ 9.9 63.5 ⫾ 7.2

1 Balanced diet concerning all minerals and vitamins; one puppy was removed from the group for other then trial reasons.
* Significant differences between the three trials in the control group were determined by Tukey’s test (P ⬍ 0.05). Letters identify differences
between more than two groups; means not sharing a superscript letter are significantly different (Tukey’s test, P ⬍ 0.05).
 DIGESTIBILITY OF CA AND P IN GROWING DOGS
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