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ANIMAL GROWTH, PHYSIOLOGY, AND REPRODUCTION |



* Department of Companion Animal Clinical Sciences, Norwegian School of Veterinary Science, PO Box 8146 Dep., N-0033 Oslo, Norway;
and
Norwegian Kennel Club, PO Box 163 Bryn, N-0611 Oslo, Norway;
and
Department of Basic Sciences and Aquatic Medicine, Norwegian School of Veterinary Science; and
and
Department of Production Animal Clinical Sciences, Norwegian School of Veterinary Science
| Abstract |
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Key Words: alkaline phosphatase body weight canine growth curve normal development puppy
| INTRODUCTION |
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Mathematical models of growth functions can be used to summarize growth data for an individual or a population. The Gompertz function was introduced for this purpose (Winsor, 1932
) and has proven useful for estimating growth curves for dogs (Allard et al., 1988
; Helmink et al., 2000
) and various other species (Menchaca et al., 1996
; Emmans and Kyriazakis, 1997
; Topal et al., 2004
).
During growth, the metaphyses of the long bones remodel continuously to create the mature skeleton (Kincaid and Van Sickle, 1983
). In a study by Voorhout et al. (1994)
, the length of the radius and ulna was measured in Great Danes, using radiographs. To the best of our knowledge, the circumference of the distal radius and ulna (CDRU) has not previously been described during growth.
Alkaline phosphatase (ALP) isoenzymes are frequently measured in relation to skeletal growth and disease. Three ALP isoenzymes, liver ALP, bone ALP (BALP), and corticosteroid-induced ALP are present in the dog (Hoffmann and Dorner, 1975
; Saini et al., 1978
; Mahaffey and Merrill, 1991
). Bone ALP is synthesized by, and expressed on, the external surface of the osteoblast. Allen et al. (2000)
reported that BALP provided the main contribution to the total ALP activity in dogs less than 1 yr of age.
The principal objective of the current study was to describe growth patterns in 4 dog breeds raised in Norwegian domestic environments and the concomitant changes in 2 growth-related clinical variables, total ALP and CDRU. The second objective was to investigate how independent variables like age, sex, litter number, and birth weight affected these clinical variables.
| MATERIALS AND METHODS |
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Study Design
A longitudinal observational study was conducted to study growth variables in 4 large breeds of dog: New-foundland (NF), Labrador retriever (LR), Leonberger (LEO), and Irish wolfhound (IW). The present paper is a part of a larger study designed to study environmental factors influencing growth and the occurrence of skeletal diseases.
Seven hundred privately owned dogs derived from 107 litters were initially enrolled in the study (each litter was assigned a number as identification, which was designated as litter number), each one with a housing and feeding regimen decided by its owner.
Inclusion of Animals
Eligibility for the study was that the dogs of the selected breeds were born in Norway between November 1998 and June 2001. The breeding stock consisted of dogs born in Norway and dogs that had been imported. Each breeder, dog owner, and veterinarian that participated in the project signed a written agreement of cooperation to comply with the project plan. Inclusion of a litter in the project began when the bitch was mated. The puppies were registered in the Norwegian Kennel Club. Not all animals initially enrolled in the study continued to completion (Table 1
). Reasons for non-completion included but were not limited to death of the dog, relocation of the owners during the study, and misunderstanding that the study continued after the hip and elbow radiographs. Additionally, some dogs missed one or more of the examinations during the study for unknown reasons.
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The breeder was asked to record the BW of each puppy at birth and on d 3 and 7, and then weekly until 8 wk. Feeding regimens of each litter were decided by the breeder. All puppies stayed with their mother from birth to approximately 8 wk, when they were delivered to the new owner. The new owners completed the questionnaires and reported information at the observational ages of 3, 4, 6, 12, 18, and 24 mo. The numbers of completed questionnaires received are given in Table 2
.
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Laboratory Analysis
Hematological tests were conducted, and the blood serum was analyzed for 22 biochemical variables, including total ALP. Total ALP was analyzed with the diethanolamine (DEA) method, recommended by the Scandinavian Committee on Enzymes. In the DEA method, the sample is added to a p-nitrophenyl phosphate substrate, and DEA buffer is used to maintain the reaction pH at 9.7 to 9.8. Magnesium ions are added to the DEA buffer to activate and stabilize the enzyme. During the reaction, ALP hydrolyzes the p-nitrophenyl phosphate to form p-nitrophenol, which can be measured photometrically at 405 nm. The reaction rate follows zero-order kinetics. The units of activity can be calculated as micromoles of substrate hydrolyzed per minute, which is based on the molar absorptivity of p-nitrophenol. The analyses were conducted at Sentrallaboratoriet, Norwegian School of Veterinary Science, Oslo, Norway.
Statistical Analysis
Body weight data separated by breed and sex were fitted to a Gompertz function. A Gompertz function is a nonlinear, sigmoid function with its point of inflection at 36.8% of mature BW. Growth was modeled with the following equation (Helmink et al., 2000
), using the NLIN procedure (SAS Inst. Inc., Cary, NC):
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where Wt = BW at time t, Wmax = mature BW, b = constant proportional to duration of growth, c = age at point of inflection (age at which 36.8% of mature BW is reached), and t = age in days.
Analyses were carried out separately for each breed and sex. Duration of growth was estimated as 4b + c, which describes 98% of the growth phase (Helmink et al., 2000
). The derivative of the Gompertz function describes the growth rate.
Longitudinal modeling assessed the relationship between BW and various potential predictors of BW using the MIXED procedure of SAS. Log-transformed BW values were used, as they provided a better fit to the normal distribution than the original values. Analyses of the independent variables (log BW, total ALP, and CDRU) were performed on the total data set as well as within breed. Age and sex were included as explanatory variables. Other explanatory variables (litter number, birth weight) were included when significant for at least one of the breeds. Interactions were evaluated by the likelihood ratio test from models with and without the interaction terms. When assessments were made of the relationships between the change of BW or change of total ALP and potential predictors, transformation of the data was unnecessary.
| RESULTS |
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Weight gain, expressed by the derivative of the Gompertz function, reached its maximum value between 89 and 106 d (point of inflection), after which it gradually declined as mature BW was achieved (Figure 1
). The data indicate that the peak BW gain in females occurred on average 6 d earlier than in males and that their overall BW gain was less than in males after the greatest growth period.
Circumference of the Distal Radius and Ulna
The estimated average CDRU increased after 90 d and peaked at 180 d. Thereafter, CDRU declined and stabilized at about 1 yr of age (Figure 2
). At comparable ages, males had a greater (P < 0.001) CDRU than females. In mixed-model analyses of the whole data set, CDRU was affected (P < 0.001) by age, breed, sex, and to the interactions age x breed and age x sex (P = 0.04). Differences between sexes and among breeds increased with age. Within breeds, CDRU was affected by age and sex. The effect of litter number on CDRU was significant in all breeds (P < 0.001) except IW (P = 0.6). Birth weight showed a positive relationship with CDRU (P < 0.001) in LR. In NF, there was a trend (P = 0.07) for a positive relationship, whereas in the other breeds there was no significant relationship (P = 0.2 for LEO and P = 0.6 for IW).
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| DISCUSSION |
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More than 33% of adult BW was reached at approximately 3 mo of age, more than 1 mo earlier than suggested by Meyer and Zentek (2001)
. Their recommendation was that puppy BW should not exceed 33% of adult BW after the fourth month, 60% after the sixth month, and 80% after 12 mo. Our data showed that these suggestions were exceeded in all breeds at all ages (Figure 1
). A possible explanation for the apparent discrepancy between our results and previously reported growth data for similar breeds is that our study included privately owned, healthy dogs. Much of our present knowledge originates from experimental studies, such as feeding experiments, which might not be an accurate reflection of life for family dogs (Hedhammer et al., 1974
; Hazewinkel et al., 1985
; Kealy et al., 1997
).
Only 2 previous studies (conducted in Switzerland and the United States, respectively) have reported growth patterns in dogs based on the Gompertz function. Helmink et al. (2000)
studied LR and reported mature female BW at 319 d (26.8 kg) and mature male BW at 335 d (31.4 kg). Allard et al. (1988)
found very similar results. Both these studies differ from the current study by reporting a faster growth pattern and a lower mature BW for both sexes. It is worth noting that Helmink et al. (2000)
studied a population that has been subject to several generations of genetic selection for ability to perform in a specific working role, and the dogs need to fall within a narrow range of body sizes to work effectively. It is possible that there has been a correlated response in growth rate for selection as suitability to work as guide dogs. In a study on BW changes during growth of 12 breeds, including NF, IW, and LR, (Hawthorne et al., 2004
), sex effects were not included. In their study, NF dogs (7 males and 8 females) reached mature BW at 46 wk of age, approximately 12 wk earlier than dogs in the current study. The results for IW (2 males and 5 females) were similar to those for NF, but for LR (16 males and 21 females) the estimates were comparable with our findings. Differences between the results in this study and the results in the study of Hawthorne et al. (2004)
may be attributed to sample-size effect. Hawthorne et al. (2004)
concluded that as well as differences in growth patterns among toy, small, and large breeds, there were additional differences in growth patterns among breeds of similar size.
The effects of age, breed, sex, birth weight, and litter number were significant predictors of log BW in our study. Although sex was not significant for IW, this was probably due to the low number of observations for this breed. Growth curves for various dog breeds have often been presented with both sexes in a single curve (Kealy et al., 1992
, 1997
; Hawthorne et al., 2004
). Our findings suggest that growth curves for dogs should be presented separately for breed and sex, as also previously suggested by others (Allard et al., 1988
; Helmink et al., 2000
).
Allard et al. (1988)
, studying Beagles, Brittany spaniels, and LR, reported that the BW difference between males and females of a breed was insignificant at birth. In the study by Allard et al. (1988)
, birth weight was 490 g in females and 480 g in the male LR. These are considerably heavier than those found in our study, despite mature BW in our study being heavier than those reported by Allard et al. (1988)
. Our study also indicated that birth weight was influenced by sex in at least 3 of the breeds studied (LR, LEO, and NF).
To the best of our knowledge, no other reports have described the litter effect on BW and growth in dogs. The litter effect mostly expresses the maternal effect on growth but is highly complex because it encompasses environmental and genetic factors. In a study of swine, maternal genetic effects were tested in 4 breeds. The indication was that litter effect, environmental, and maternal effects were important for 100-d BW in all 4 breeds (Johnson et al., 2002
). The litter effect should be studied more thoroughly to elucidate how maternal and environmental factors contribute to growth and development in dogs.
The CDRU had already reached maximum size at 6 mo of age, with the greatest circumference in our study found in NF dogs. Although the relatively early development is interesting, from a clinical point of view, the significance of this is uncertain. The CDRU is a readily available variable for veterinary clinicians to measure, and it is valuable to have normal reference values to enable pathological metaphyses to be distinguished. In dogs with metaphyseal osteopathy, the metaphyses of radius and ulna swell (Grondalen, 1976
).
To our knowledge, this is the first study that has compared total ALP in large, rapidly growing breeds and investigated its variation with respect to explanatory variables like litter number, sex, and BW change. The general trend was a decrease in total ALP with increasing age and that breed differences diminished with increasing age. These results indicate that age and breed are the dominant explanatory variables and that BW change in the period between 4 and 6 mo of age explained a relatively small proportion of the variation in the change in total ALP during the same period, although a significant positive relationship was found. The analyses of total ALP indicated that although significant differences were found among breeds, these differences were not due to differences in body size; LR had the second greatest total ALP concentration at 90 d and was the smallest breed in the study. The expectation would be that large dogs would have the greatest total ALP concentrations. Surprisingly, LR had a greater concentration of total ALP than the NF and the LEO from 90 to 180 d of age. In a study of markers of bone metabolism in growing LR, Breur et al. (2001)
concluded that BALP is a moderately strong indicator of bone length growth. In skeletally immature animals, the activity of BALP will tend to predominate because of high basal bone turnover rates in growing animals, and a strong correlation between total ALP and BALP has been reported in dogs less than 1 yr old (Allen et al., 2000
). In another study, Allen et al. (1998)
showed that activities of total ALP and BALP followed similar trends.
In a study on markers of bone metabolism in dog breeds of different size, there was no significant difference in the concentrations of BALP and type I collagen in the adult Pomeranian and IW (Breur et al., 2004
). The findings in their study support our finding that the differences in total ALP among breeds disappear with increasing age.
Allen et al. (1998)
gave a mean reference value for total ALP of 100.2 (±20.9) for dogs (n = 10) in the age group 0 to 1 yr. In our study, the concentration of ALP differed among breeds but decreased in all breeds from 90 to 360 d of age. Age was the major explanatory variable of the change in total ALP concentration during growth. This emphasizes the importance of estimating the concentration of this enzyme at different ages during growth. In an experimental study on the influence of chronic calcium excess in Great Danes during growth, total ALP was measured (Hazewinkel et al., 1985
). The concentration in the calcium-exposed group (n = 6) had very similar values to the IW in our study. However, the control group (n = 5) did not show any decrease in total ALP with increasing age. A possible explanation for the apparent discrepancy between our results and those of Hazewinkel et al. (1985)
may be sample-size effect.
| IMPLICATIONS |
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| Footnotes |
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2 Corresponding author: cathrine.trangerud{at}veths.no
Received for publication June 2, 2006. Accepted for publication August 11, 2006.
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