WALTHAM International Science Symposium:
Nature, Nurture, and the Case for Nutrition
Urinary Composition of Cats Is Affected by the Source of Dietary Protein1
Jürgen Zentek2,3 and Annette Schulz
Institute of Animal Nutrition, School of Veterinary Medicine Hannover, D30173 Hannover, Germany
EXPANDED ABSTRACT
KEY WORDS:
cats
urolithiasis
urinary excretion
oxalate
crystals
Feline lower urinary tract disease is an important clinical
problem in cats. Epidemiological data based on urinary stones
submitted for examination show over the last decade
a consistent trend for struvite calculi to decrease and calcium
oxalate to increase (1,2). The risk for cats to develop
urolithiasis is related to dietary and nondietary factors.
Higher solute concentrations, a subsequent supersaturation of
the urine, and the lower frequency of urination can favor the
formation of crystals and calculi. Although urinary calcium
excretion is normally low in cats and not strongly related to
dietary mineral intake, the excretion of phosphorus, magne-
sium, sodium, and potassium was repeatedly demonstrated to
be strongly diet dependent (3). The dietary mineral composi-
tion influences the postprandial pH significantly and can
therefore predispose cats and dogs to develop crystaluria or
urolithiasis (4,5). Oxalate is important due to its potential to
form crystals with calcium. Urinary oxalate excretion is
resultant from dietary oxalate and the metabolism of ascorbic
acid, glycine, tryptophane, phenylalanine, and hydroxyproline
(6). Experimentally, a vitamin B-6 deficiency was found to
increase the daily urinary oxalate excretion from 1.5 to 24 mg
BW1 (7). Similar observations were made in young cats, fed
either a vitamin B-6–deficient diet or a diet supplemented with
8 mg/kg vitamin B-6 (8,9).
Dietary protein intake could be an important factor
determining the urinary composition in cats, but has not been
intensively studied. Urine is the most important excretory
medium for metabolites from protein metabolism. Urine
volume shows a positive correlation with dietary protein intake
(3). Urea, creatinine, and ammonia are the main metabolites
that are excreted. Even in healthy cats a certain level of protein
excretion is considered as normal.
1
Presented as part of the WALTHAM International Science Symposium:
Nature, Nurture, and the Case for Nutrition held in Bangkok, Thailand, October
28–31, 2003. This symposium and the publication of the symposium proceedings
were sponsored by the WALTHAM Centre for Pet Nutrition, a division of Mars, Inc.
Symposium proceedings were published as a supplement to The Journal of
Nutrition. Guest editors for this supplement were D’Ann Finley, James G. Morris,
and Quinton R. Rogers, University of California, Davis.
2
To whom correspondence should be addressed. E-mail: juergen.zentek@
vu-wien.ac.at.
3 5
Present address is Institute of Nutrition, Veterinary University of Vienna,
Vienna, Austria.
0022-3166/04 $8.00 Ó 2004 American Society for Nutritional Sciences. J. Nutr. 134: 2162S–2165S, 2004.
2162S
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In this study the influence of dietary protein quality and
quantity was investigated with regards to the urine composition
and the excretion of oxalate and crystaluria in cats.
MATERIAL AND METHODS
Six diets were formulated to contain either collagen tissue, soybean
protein isolate, or horsemeat as protein sources. Each protein source
was tested in two concentrations to give a low- or a high-protein
variant (see Table 1 for ingredients). For both soy diets it was
necessary to add 5% of porcine liver to improve palatability.
Collagen tissue (CT)4 (Fettschmelze, Versmold, Germany), soy
protein isolate (SI) (Sojamin 90, Lucas Meyer GmbH, Hamburg,
Germany), and horsemeat (HM) (Jausch, Pattensen, Germany) were
used as protein ingredients. Dry matter of the ingredients was 96.5,
94.1, and 27.7%; crude protein was 76.2, 82.6, and 20.8%; and crude
fat was 13.3, 4.3, and 4.6% (all on a fresh matter basis).
The nutrient and mineral composition of the diets is given in Table
2. The dry diets with collagen tissue and soy protein were mixed with
measured amounts of distilled water to improve palatability.
The diets were fed sequentially to seven adult cats (five females,
two males, mean age 4 y, body weights 3.99 6 0.76 kg)5. The diets
were fed consecutively (collagen tissue, soy protein isolate, horsemeat)
with the high-protein diet followed by the low-protein diet. Each
feeding period lasted 28 d and the cats were fed a commercial mixed
diet for 28 d between each change of the dietary protein source. During
the adaptation period the cats were kept in groups, for the collection of
feces and urine individually in metabolism cages. Each diet was fed for
at least 14 d before samples were collected. The food amounts were
calculated individually to meet the maintenance requirements and to
keep the bodyweights constant. Food was offered once daily in the
morning. The following analyses were carried out: Weende (10) and
mineral analysis of food, urinary excretion of nitrogen, urea, ammonia,
protein, oxalate and creatinine, urine pH, urine specific gravity, and
number and size of urinary crystals. Minerals were determined after
wet washing in a mixture of perchloric and nitric acid. Atomic
absorption spectrophotometry was used to study calcium and
4
Abbreviations used: CTHP, collagen tissue, high-protein diet; CTLP, collagen
tissue, low-protein diet; DM, dry matter; HMHP, horsemeat, high-protein diet; HMLP,
horsemeat, low-protein diet; ns, not significant; SD, standard deviation; SIHP, soy
protein, isolated, high-protein diet; SILP, soy protein, low-protein diet.
Care and treatment of the animals was approved by the governmental
committee according to the German Tierschutzgesetz.
 PROTEIN INTAKE AND URINE COMPOSITION IN CATS 2163S

TABLE 1
Ingredients of the experimental diets

Protein Animal Porcine Vitamin/mineral

Protein source Diet1 source Rice fat Cellulose liver supplement2

%
Collagen tissue3 CTHP 85 9 — 3 — 3
CTLP 25 50 19 3 — 3
Soy protein isolate SIHP 70 9 10 3 5 3
SILP 25 45 19 3 5 3
Horsemeat HMHP 95.5 2.8 — 0.9 — 0.9
HMLP 55 30 11.3 1.9 — 1.8

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1 Diets designated according to the protein source: CT
HP collagen tissue, high protein; CTLP collagen tissue, low protein; SIHP soy protein, isolated,
high protein; SILP soy protein, low protein; HMHP horsemeat, high protein; HMLP horsemeat, low protein
2 Vitakalk (MFE Marienfelde GmbH, Roth, Germany), composition: 21% calcium, 6% sodium, 8% phosphorus, 1% magnesium, 300 mg/kg copper,
3000 mg/kg zinc, 350 mg/kg manganese, 2500 mg/kg iron, 80 mg/kg iodine, 15 mg/kg cobalt, 5 mg/kg selenium, 250,000 IU/kg vitamin A, 10,000 IU/kg
1,25-dihydroxycholecalciferol, 1000 mg/kg vitamin E, 50 mg/kg thiamin, 100 mg/kg riboflavin, 80 mg/kg pyridoxine-HCl, 1000 mg/kg cyanocobalamin,
1000 mg/kg vitamin C, 15 mg/kg menadione, 1000 mg/kg nicotinic acid, 500 mg/kg pantothenic acid, 5000 mg/kg biotin, 30 mg/kg folic acid, and 5000
mg/kg choline chloride.
3 CT, dried protein, remaining after fat melting from porcine and bovine tissues, consisting mainly of connective tissue.

magnesium (11), flame photometry for sodium and potassium (12), subjectively graded depending on the size (large .150 mm, in-
and photometry (vanadate molybdate method) for phosphorus (13). termediate 50–150 mm, small ,50 mm). Microscopy was performed
Deionized drinking water was offered ad libitum and daily water daily over 4 d and averages were calculated per cat. Creatinine was
consumption was recorded after correction for evaporative losses. measured by a modified Jaffé method (Nobi Flow, Kreatinin-JE, Nobis,
Urine volume was measured in two 4-d periods in metabolism cages. Endingen, Germany) and urea by urease method (NobiFlow,
Urine was collected in acidified containers (addition of 10 mL 1 N Harnstoff-Rapid, Nobis, Endingen, Germany). Oxalate concentrations
hydrochloric acid, Merck AG, Darmstadt, Germany) during the first were determined by an enzymatic method (Sigma oxalate test, Sigma,
4 d for the analysis of nitrogen, urea, ammonia, oxalate, and creatinine Deisenhofen, Germany). Protein concentrations were measured
and in nonacidified containers during the last 4 d for the analysis of from fresh urine samples by the Coomassie brilliant blue–G-250
protein, pH, specific gravity, and sediment. Thymol (Fluka Chemie method (15).
GmbH, Buchs, Switzerland) was added to prevent bacterial growth
(14). Urine production was monitored by regular inspections of the Statistics
cages over the day (every 2 h between 0800 and 1800 h) and
overnight. Urine volume was determined by a scaled measuring Data were processed by EXCEL 5.0 and SAS 6.04 (16). Data are
cylinder with 1-mL graduations and corrected for the added summarized as means 6 SD. After testing for homogeneity, two-factor
hydrochloric acid. Urine samples from each cat were divided into ANOVA (factors protein quality and quantity) was performed. In case
aliquots and investigated freshly or frozen at 208C until analysis. of significant interaction of the main factors the feeding periods were
Fresh samples were used for determination of specific gravity with compared by t test. P-values ,0.05 were taken as significant.
a refractometer (Krüss GmbH, Hamburg, Germany) after temperature Regression analysis was performed to determine the correlations
compensation. Urinary pH was measured in fresh samples by a digital between different traits.
pH meter (WTW, Model pH 526, Weilheim, Germany), and ammonia
in the acidified urine samples after thorough homogenization was
measured by an ammonia-sensitive electrode (Philips, Model IS570 RESULTS
NH3, Kassel, Germany) that was adapted to a digital mV-Meter
(Knick, Berlin, Germany). Urine sediment was investigated in fresh All cats ingested the experimental diets voluntarily.
samples by light microscopy after 10 min of centrifugation at 1500 3 g Palatability was good for diets HMHP and SILP but was reduced
(Biofuge, Christ, Osterode, Germany). One drop of the sediment was for all other diets that were ingested only slowly. Feed
investigated in a microscope by 100-fold magnification. Ten fields were
consumption was 11.0–15.6 g  kg of BW1d1 (CTHP 12.6
counted for the numbers of crystals and cells. Crystals were
6 4.2, CTLP 13.8 6 4.9, SIHP 11.4 6 3.1, SILP 15.6 6 2.9,
HMHP 15.0 6 2.2, HMLP 11.0 6 3.1) resulting in an
metabolizable energy intake of 244–330 kJ  kg of BW1d1
TABLE 2 (CTHP 264 6 88, CTLP 310 6 111, SIHP 244 6 66, SILP 349 6
Nutrient and mineral analysis of the experimental diets (% DM) 64, HMHP 330 6 48, and HMLP 245 6 68). Body weights were
kept constant in the collection periods in a range from 160 to
CTHP CTLP SIHP SILP HMHP HMLP 1100 g/8 d. With the soya diets three of the seven cats had soft
feces, occasionally with visible traces of blood and mucus. The
Crude ash 4.73 3.50 5.43 3.68 5.37 3.41 urine volume (y) was determined mainly by total water intake
Crude protein 77.6 22.64 64.2 27.3 64.0 21.7 (y ¼ 0.55x 1 0.28; r2 ¼ 0.65; P , 0.001) (Table 3). The cats
Crude fat 10.1 23.5 14.7 21.8 16.9 22.2 consumed 18.1–44.0 mL water  kg of BW1d1 by feed and
Crude fiber 2.78 2.90 2.61 3.02 2.68 2.17
Nitrogen-free extract 4.79 47.5 13.1 44.2 11.1 50.6 drinking water. The renal water excretion was low with the diet
Calcium 1.00 0.86 0.76 0.87 0.79 0.71 HMLP (8.3 mL  kg of BW1d1) and was 14.6–26.4 mL  kg of
Magnesium 0.08 0.06 0.09 0.07 0.10 0.07 BW1d1 in the other dietary periods.
Phosphorus 0.76 0.46 0.62 0.37 0.60 0.49 Urinary pH (Table 4) was lower with the low-protein diets,
Chloride 0.99 0.57 0.69 0.44 0.54 0.36 except when horsemeat was fed. The specific gravity of urine
Sodium 0.81 0.37 1.29 0.50 0.38 0.26 was influenced by the urine volume and dietary protein source
Potassium 0.67 0.22 0.15 0.15 1.22 0.33
and quantity. The nitrogen intake explained a small percentage
2164S SUPPLEMENT

TABLE 3 linearly correlated to the urinary concentration of oxalic acid

(y ¼ 0.83 3 e0.0016, r2 ¼ 0.47; P , 0.05) and also with
Total water consumption and fecal and urinary water the urinary excretion of oxalate (y ¼ 6.50 3 e0.0009, r2 ¼ 0.24;
excretion in the experimental cats P , 0.05). However, calcium oxalate dihydrate crystals were
observed only in one urine sample during the period HMHP.
Total Fecal Urinary The number of struvite crystals (y) was considerably higher
water water water (Table 5) and depended on the quantity and quality of dietary
consumption excretion excretion
protein but was also related to urinary pH (y ¼ 6.36x  39.2,
Diet mL  kg of BW1d1 r2 ¼ 0.24; P , 0.001).
CTHP 44.0a 6 13.1 3.66ab 6 1.56 21.4a 6 7.45
CTLP 30.1b 6 10.7 3.46abc 6 2.96 14.6b 6 6.37
SIHP 34.3ab 6 8.98 2.52b 6 1.41 21.6a 6 6.86
SILP 36.3b 6 6.73 4.64a 6 3.04 18.2ab 6 3.59 DISCUSSION
HMHP 35.3b 6 6.25 0.97c 6 0.41 26.4a 6 5.16

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HMLP 18.1c 6 4.89 1.38bc 6 0.84 8.3c
Means within a column not sharing a common superscript are
significantly different at P , 0.05 (ANOVA and post-hoc t test). n ¼ 7,
means 6 SD.
of the variation of urine specific gravity (r2 ¼ 0.20; P , 0.01)
but the urinary concentration of urea was strongly (r2 ¼ 0.83;
P , 0.001) correlated to the specific gravity of urine. Urinary
nitrogen excretion was 399–553 mg  kg of BW1d1 with the
low-protein diets and increased to 960–1426 mg  kg of
BW1d1 with the high-protein diets. Urea excretion (y) was
1086–1488 mg  kg of BW1d1 with the low-protein diets and
increased to 2557–3672 mg  kg of BW1d1 with the high-
protein diets in strong correlation to N intake (y ¼ 1.82x 1
389, r2 ¼ 0.80; P , 0.001).
Urinary ammonia excretion (y) and the relation to nitrogen
intake (x) was more variable compared to urinary nitrogen and
urea excretion. It increased significantly with nitrogen intake
(y ¼ 0.022x 1 3.56, r2 ¼ 0.59; P , 0.001) and was also
influenced by the dietary protein source. Urinary creatinine
(y ¼ 0.0078x 1 30.19, r2 ¼ 0.22; P , 0.01) and protein
excretions (y ¼ 0.0019x 1 2.23, r2 ¼ 0.45; P , 0.001) were less
strongly correlated to dietary nitrogen intake (x). Dietary
protein source affected only creatinine excretion.
The dietary protein quantity and quality affected urinary
oxalate excretion. Nitrogen intake (x) was inversely and not
6 1.91 The study demonstrates that dietary protein intake in-
fluences urine composition and crystaluria in cats. The protein
sources used in the experimental diets had different qualities.
Collagen tissue has a lower biological value compared to meat
with lower concentrations of most essential amino acids and
high levels of hydroxyproline and glycine (3). Compared to
meat as dietary protein the total digestibility is reduced in cats
(3) and at the ileal level in dogs (17). Horsemeat was chosen as
a high-quality protein source and also because it could be
expected that the experimental cats were not exposed to this
protein before. Soy protein isolate was chosen as a vegetarian
protein source. Due to the low palatability 5% porcine liver had
to be included in the soy diets. On a dry matter basis, the added
amount of protein is low compared to the fraction from soy, but
this is confounding the experimental design. However, it
became obvious, that dietary protein intake is an important
determinant of urine composition in cats. Protein intake
correlated positively with urine specific gravity, nitrogen, and
urea excretion, and less strongly also with the creatinine and
ammonia excretion. Urinary ammonia excretion depends on
the dietary protein intake but is also involved in the regulation
of acid base metabolism. A lower urinary pH (x) was associated
with higher ammonia concentrations (y, mg/mL) (y ¼ 3.79 
0.34x; r2 ¼ 0.11; P , 0.05). Urinary creatinine was influ-
enced by protein intake and additionally by protein quality. The
highest creatinine excretions were found with the collagen
tissue diets, whereas the horsemeat diets were intermediary and
soy protein induced the lowest urinary creatinine excretion.

TABLE 4
Urine pH, specific gravity, and urinary excretion of nitrogen, urea, ammonia, creatinine, protein,
and oxalic acid in the experimental cats

Nitrogen Urea Ammonia Creatinine Protein Oxalic acid

Diet pH Specific gravity Urinary excretion, mg  kg of BW1d1

CTHP 7.43b 6 0.35 1.056a 6 0.006 1065b 6 352 2870b 6 916 29.5b 6 6.7 50.7 6 12.0 4.93 6 1.36 2.83c 6 0.89
CTLP 6.70c 6 0.19 1.039c 6 0.006 440d 6 149 1301cd 6 417 15.3d 6 3.8 38.8 6 7.9 2.38 6 0.93 13.70a 6 4.32
SIHP 8.26a 6 0.31 1.051b 6 0.007 960b 6 283 2557b 6 960 21.6c 6 7.1 33.2 6 3.7 5.02 6 2.14 1.17d 6 0.53
SILP 7.56b 6 0.31 1.034d 6 0.004 553c 6 121 1488c 6 365 14.6d 6 4.2 28.5 6 2.6 4.18 6 1.12 4.79b 6 2.73
HMHP 7.10b 6 0.12 1.056a 6 0.003 1426a 6 210 3672a 6 556 55.2a 6 10.6 39.1 6 3.3 4.64 6 1.27 0.94d 6 0.29
HMLP 7.15b 6 0.45 1.050b 6 0.010 399d 6 58 1086d 6 240 14.2d 6 2.8 36.6 6 4.9 3.46 6 0.86 3.62b 6 2.44
Two-factor ANOVA (P-values)
Dietary protein 0.001 0.01 ns ns 0.001 0.01 ns 0.001
source
Dietary protein 0.001 0.001 0.001 0.001 0.001 0.01 0.01 0.001
quantity
Source x 0.01 0.05 0.01 0.05 0.001 ns ns 0.001
quantity

Means within a column not sharing a common superscript are significantly different at P , 0.05, two-factor ANOVA, and t test for group comparison in
case of significant interaction of protein source and quantity. n ¼ 7, means 6 SD.
 PROTEIN INTAKE AND URINE COMPOSITION IN CATS 2165S

TABLE 5 Conclusion
Number of small, large, and large struvite crystals per Dietary protein intake and the source of protein determined
microscopic view area (100-fold magnification of urinary excretion of nitrogen metabolites and oxalate and the
urine sediment) in the experimental cats
level and character of crystaluria. The significance of protein
intake should be further investigated with regards to feline
Struvite crystals lower urinary tract disease.
Diet Small Medium Large

CTHP 2.06 6 2.30 0.49 6 0.34 0.09 6 0.07

CTLP 0.48 6 0.53 0.13 6 0.03 0.10 6 0.09
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