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* Facultad de Ciencias Agrarias, Universidad Nacional del Litoral, P. Kreder 2805, Esperanza 3080, Argentina
Universidade de Passo Fundo, Faculdade de Agronomia e Medicina Vetorinaria, Barrio São Jose, Brazil
Department of Animal Science, McGill University, Ste-Anne-de-Bellevue, Quebec H9X 3V9, Canada
Corresponding author: R. I. Cue; e-mail: Roger.Cue{at}McGill.ca.
| ABSTRACT |
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Key Words: functional herd life Holstein survival analysis type trait
Abbreviation key: FHL = functional herd life, LPL = length of productive life, RCR = relative culling rate, PATLQ = Programme dAnalyse des Troupeaux Laitiers du Quebec
| INTRODUCTION |
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However, longevity is a difficult trait to measure, mainly due to the presence of incomplete records (i.e., cows that are still alive when the study is carried out). Ducrocq (1987) defined functional herd life (FHL) as the ability of a cow to remain sound and healthy after adjustment for production level; it refers to the ability to delay involuntary culling. Correcting the length of productive life of a cow for phenotypic milk production is used as an approximation for FHL.
Survival analysis, which deals with censoring in the analysis of time response, has been applied in the animal breeding context to estimate genetic parameters for longevity. It is the methodology applied for routine genetic evaluation for longevity in dairy cattle in France, The Netherlands, Germany, Denmark, The Czech Republic, and Italy (Van der Linde and de Jong, 2003). Length of productive life (LPL), defined as the number of days from first calving to death, culling, or censoring (Ducrocq et al., 1988), is a suitable measure because it is characterized by the presence of incomplete records.
The objective of this study was to evaluate the effects of conformation traits on FHL by means of survival analysis.
| MATERIALS AND METHODS |
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The data were merged with 23 conformation traits evaluated by the Holstein Association of Canada; 58% of the records had type information, from which 153,479 (80.3%) corresponded to the official option and 37,688 (19.7%) to the owner sampler option. Therefore, most of the type information corresponds to cows in the official herds. Type information consisted of actual phenotypic type scores of 15 linear descriptive traits (evaluated on a 9-point scale) and 6 composite traits (evaluated on an 18-point scale). Composite traits are calculated directly from the linear score traits (weighting score for each component trait, Holstein Canada, 2003). Front end, body depth, rear attachment width, udder depth, fore teat length, and dairy form linear traits were not included because of limited data. Table 1
shows the 15 descriptive linear traits included in the analysis, the method of evaluation, a brief description, and the optimal score for each (Holstein Canada, 2003). For all of the composite traits, a high score is desirable (Holstein Canada, 2003). When a cow is evaluated, the classifiers hand-held computer uses the name, birth date, calving date, lactation number, and a rating for the fullness of the udder (whether the cow has been recently milked or not) to calculate adjustment factors for the mentioned effects when converting the measurements to a descriptive linear code scored from 1 to 9 for on-farm use. The type information corresponded to first classifications. Information about the pedigree status of the cows and about classification (stage of lactation at classification, herd-classifiers, etc.) were not available.
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Model
The following Weibull model was used:
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where
(t) is the hazard function of a cow t days after calving;
o (t) [
(
t)
- 1] is the Weibull baseline hazard function, with scale parameter
and shape parameter
(the parameter
was fixed at 1.8, which was estimated in a previous analysis); yi (t') is the time-dependent effect of year of calving, with changes on March 1 of each year; pj (
) is the time-dependent effect of lactation number x stage of lactation (lactation 1, 2, 3, and 4 x 4 stages of lactation), with changes occurring at
= 0, 120, 240, and 305 d of each lactation; zk (t') is the time-dependent effect of annual change in herd size k. Seven classes were defined: a decrease >25%, a decrease of 15 to 25%, a decrease of 5 to 15%, no appreciable change (-5 to +5%), an increase of 5 to 15%, an increase of 15 to 25%, and an increase >25%, with changes at March 1 of each year; am is the time-independent effect of age at first calving (22 to 40 mo of age); du is a dummy variable that indicates the presence or absence of type information. This variable was included to allow all the cows to be considered in the analysis, even the ones without type data, in order to avoid bias. hn (t') is the random time-dependent effect of herd-year, with changes at t' = March 1 of each year (herd-year was assumed to follow a log-gamma distribution, which was algebraically integrated out from the joint posterior density); typeb is the time-independent effect of type trait; and wr (
) is the time-dependent effect of within herd-year-parity class of milk production at 305 d of lactation (first or later lactation). Five classes were considered: <1.5 SD, between -1.5 and -0.5 SD, between -0.5 and +0.5 SD, between +0.5 and +1.5 SD, and >1.5 SD, with changes at the beginning of a new lactation. The 305-d yields for all cows were standardized to 4% fat and 3.3% protein (PATLQ). The following composite type traits were analyzed: final score, frame-capacity, rump, feet and legs, mammary system, and dairy character. The 15 linear type traits analyzed were stature, size, chest width, loin strength, pin setting, pin width, foot angle, bone quality, rear leg set, median suspensory, udder texture, fore attachment, front teat placement, rear attachment height, and rear teat placement.
Phenotypic scores were considered as class effects. No specific relationship between type traits and functional herd life was assumed. Analyses were performed for each type trait one at a time; hence, 21 separate analyses were performed on the same dataset according to the model previously described. The Survival Kit V3.0 by Ducrocq and Sölkner (1998) was used. The overall influence of each type trait on FHL was assessed using the likelihood ratio test. Significance was tested by adding each effect in sequential order in the model.
| RESULTS |
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Figures 1
and 2
show the contribution of composite and linear type traits in the change of the log likelihood (-2log L), respectively. All of the traits were significant at P < 0.001 level. Among the composite traits, the strongest effects were found for final score and mammary system, followed by feet and legs and rump. The linear traits related to the udder had the strongest effect on the FHL compared with mobility and body traits. Rear attachment height, fore attachment, udder texture, and median suspensory were the udder traits that had the strongest relationship with FHL. Among the mobility traits, bone quality had more impact on FHL than foot angle and rear legs set. Stature had the strongest effect on FHL compared with the other body traits.
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Solutions for the environmental effects indicated a higher probability of being culled: for primiparous cows calving at older ages, for cows producing below herd average, for first-lactation cows in the first or last stage of lactation, and for older cows at the end of lactation. The risk of being culled did not change for cows in herds decreasing or increasing in size.
Figure 3
shows the solutions for composite traits. The relative risks were plotted as functions of the classes of phenotypic score. All the traits followed the same trend: cows classified as poor were more likely to be culled than cows with a good classification score. The decrease in RCR was more marked for final score and mammary system compared with the other traits. For dairy character, there was 2-fold increase in risk for cows with very poor dairy character; for the last classes (14 to 18) there was a slight increase in risk, although the differences in risks were not significantly different from the reference class. For all these traits, a high score is desirable; hence, a high phenotypic score leads to an increased FHL.
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Figure 6
shows the solutions for mobility traits. Cows with coarse bones had a higher risk of being culled than a cow in the reference class; instead, the risk is reduced slightly for cows with flat and clean bones. Solutions for foot angle showed that cows with a low foot had more risk of being culled than cows in the reference class; on the other hand, cows with a steep foot had no difference in risk compared with cows with an intermediate phenotype. For this trait, the ideal score is seven (Holstein Canada, 2003). Rear leg set has an optimal score of 5 (Holstein Canada, 2003). Cows with extremely curved legs had an 80% higher risk of being culled than cows with an intermediate phenotype; on the other hand, cows with straight legs had a 34% higher risk of being culled than reference cows. This trait also displayed an intermediate optimum; however, the risk of being culled is higher for cows with extreme curved legs than for cows with extreme straight legs.
Quantification of the Effect of Mammary System on FHL
Survivor curves for cows with different phenotypes for mammary system were computed to evaluate the impact of different phenotypes on longevity (Figure 7
). For this estimation, we assumed a reference cow with average environmental conditions and a calving interval of 400 d. Three cows with different phenotypic scores were assumed: one with a good classification (class 14, favorable or desirable), one with an intermediate phenotypic score (class 10), and the last one with a poor classification (class 4, unfavorable or undesirable). Differences in survival rate for cows with different phenotypes were found. About 67% of the cows classified with a low phenotypic score, 83% of cows with an intermediate phenotype, and 90% of cows with a high phenotypic score were expected to be alive at the beginning of the second lactation (400 d after calving). Cows with a good mammary system-strongly attached, well-defined, fine-textured udders-are expected to have a longer FHL.
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| DISCUSSION |
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In the present study, dairy character showed a positive relationship with FHL. However, other studies reported a different result. Boettcher et al. (1997) found a negative genetic correlation between dairy character and herd life when it was corrected for milk production. Also, Rogers et al. (1999) found that sires that rate highest for dairy form also score low for health. It seems that refined Holstein cows with a lot of dairy character tend to be higher producers, but are more vulnerable to disease, which leads to a shorter herd life.
Among the linear traits, the strongest effects were found for udder traits. Using survival analysis, Vollema and Groen (1998), Buenger et al. (2001), Larroque and Ducrocq (2001), and Chirinos et al. (2003) reported very similar results for the Dutch, German, French, and Spanish populations, respectively. Theses studies agreed that udder traits influence the length of productive life. Boettcher et al. (1997) reported significant associations (P < 0.001) between all linear type traits and herd life for Canadian Holsteins using a linear model. Therefore, cows with well-attached udders and strong ligament are able to remain in the herd longer.
Stature and size showed a positive relationship with longevity, taller and bigger cows have better chances of surviving. Boettcher et al. (1997) reported a moderate positive relationship between body traits and herd life in Canadian Holsteins. In European populations, these traits do not contribute much to longevity (Vollema and Groen, 1998; Buenger et al., 2001; Larroque and Ducrocq, 2001). For pin setting, agreement was found with the results of Buenger et al. (2001); they reported a decrease in FHL for cows with very ascending rump. However they reported that extremely wide rump influenced negatively FHL, which is different from the results in the present study where wider rump decreased the risk of a cow of being culled.
Reasonable agreement exists between the present study and the results reported by Buenger et al. (2001) for foot angle and rear legs set; extremely low foot angles and extremely curved and straight legs led to a decreased longevity. Similar results were found by Burke and Funk (1993) for US Holsteins and Boettcher et al. (1997) for Canadian Holsteins.
As shown, there are some differences if we compare the results from Quebec Holsteins with European populations, especially for dairy character and some body traits. In our study, most of the data analyzed correspond to official herds, for which it can implicitly be accepted that official herds are interested in improving both production and conformation. These results might indicate some voluntary culling for type. It is known that in the Canadian breeding goal, conformation is an important component.
Unfortunately, we were not able to distinguish between registered and grade cows because we did not have the necessary information. For the French situation, Larroque and Ducrocq (2001) reported different relationships between conformation traits and herd life for registered and unregistered herds, especially for capacity traits. They assumed that capacity traits were subject to some voluntary culling in registered herds. Further work should look at differences between grade and registered herds to confirm if voluntary culling based on conformation exists.
Results from an earlier study using the same dataset, but including the 15 linear type traits simultaneously in the model (Schneider, 1998Schneider, 1999), showed different results. In the aforementioned study, the traits with the strongest effect on survival were rear attachment height, fore attachment, bone quality, stature, and front teat placement. Comparing the 2 studies, it is interesting to note the case of stature and size. In the present study, both traits had a similar effect and relationship with FHL, but when they were considered together in the model, size had almost no relationship with longevity. Size and stature are highly positively correlated at the genetic (0.88) and phenotypic (0.97) level (Klassen et al., 1992). The same situation was found for udder traits. When median suspensory ligament was included with the other traits, it had almost no relationship with FHL and the RCR was almost flat. Front teat placement and rear teat placement showed the same tendency. Median suspensory had high positive phenotypic (0.85, 0.96) and genetic (0.60, 0.66) correlations with front teat placement and rear teat placement, respectively. The phenotypic and genetic correlations between front teat placement and rear teat placement were (0.63) and (0.85), respectively (Klassen et al., 1992).
The results showed that the relationship between phenotypic scores and FHL is not always linear, which is in agreement with the assumptions and results reported by Larroque and Ducrocq (2001) and Buenger et al. (2001). In the present study, some traits showed an intermediate optimum, such as pin setting and rear legs set. Rear teat placement has an optimal score of 5 (Holstein Canada, 2003), but the results did not show a clear intermediate optimum, as in the case of pin setting. Buenger et al. (2001) reported that body depth, rear leg side view, and teat length exhibited an intermediate optimum. Larroque and Ducrocq (2001) did not find traits with an intermediate optimum for the French population.
Sire effects were not included in the model. It has been suggested that the effect of the type traits phenotype on culling risk could be biased because part of it is accounted for by the sire effect (Larroque and Ducrocq, 2001; Buenger et al., 2001). However, in comparing the results from the present study with a previous run (results not shown), where sire effects were included in the model, we did not find any differences in the value of the estimates nor in the significance level.
| CONCLUSIONS |
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| ACKNOWLEDGEMENTS |
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Received for publication July 17, 2003. Accepted for publication September 14, 2003.
| REFERENCES |
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