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ANIMAL GENETICS |
* USDA, ARS, US Meat Animal Research Center, Clay Center, NE;
and
USDA, ARS, US Meat Animal Research Center, Lincoln, NE
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Abstract |
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 Top Abstract INTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONIMPLICATIONSLITERATURE CITED |
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Key Words: beef cattle • breed • disease resistance • growth • heritability • shipping fever
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INTRODUCTION |
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TopAbstractINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONIMPLICATIONSLITERATURE CITED |
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; NAHMS, 2000a). In 1999, most feedlots (97.4%) within 12 states reported an overall BRD incidence of 14.4% (NAHMS, 2000a). Treatment costs for BRD averaged $15.57 per sick animal. Costs are significantly greater when labor, isolation, increased time on feed, mortality, prophylaxis, and meta-phylaxis treatments are considered (NAHMS, 2000a).
Bovine respiratory disease is a complex of diseases characterized by many types of infection, each having its own causes, clinical signs, and economic implications. Prevalent microbial causes for BRD include viral (infectious bovine rhinotracheitis, bovine viral diarrhea, bovine respiratory syncytial, and parainfluenza type 3), bacterial (Mannheimia haemolytica, Pasteurella multocida, Haemophilus somnus), and mycoplasmal (Ellis, 2001).
Shipping and processing feedlot calves enhances predisposing causes and increases environmental risk factors. Predisposing causes (Callan and Garry, 2002) are generally synergistic and include age, stress (comingling, weather, nutritional changes, etc.), and immunological background. Environmental risk factors include climate, ambient temperature, dust particles, stocking density, humidity, ventilation, and shipping distance. Epidemiological factors include microbial agent(s), mode of transmission, parasite density dependence, infectious period, latent and carrier periods, and virulence.
Heritability estimates of BRD resistance in preweaned calves have ranged from 0.10 (Muggli-Cockett et al., 1992) to 0.20 (Snowder et al., 2005). Breed types differ in incidence of BRD in preweaned calves (Snowder et al., 2005). Whether there are genetic differences among feedlot cattle is not known. Therefore, the objective of this study was to characterize environmental, genetic, and economic factors influencing incidence of BRD in feedlot beef calves.
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MATERIALS AND METHODS |
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TopAbstractINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONIMPLICATIONSLITERATURE CITED |
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) were analyzed. Cattle records spanned a 15-yr period (1987 to 2001) for pure-bred and composite breeds of beef cattle at the US Meat Animal Research Center, Clay Center, NE. Breed types included 9 pure breeds (Angus, Braunvieh, Charolais, Gelbvieh, Hereford, Limousin, Pinzgauer, Red Poll, and Simmental) and 3 composite breeds (MARC I, MARC II, and MARC III). A total of 972 sires and 6,213 dams were parents of the feedlot cattle studied. These animals were part of a germplasm use study conducted to evaluate retention of heterosis and use of breed differences in composite populations (Gregory et al., 1991, 1999). Because an upgrading breeding program had been practiced to develop a few of the purebred groups, animals with seven-eighths and fifteen-sixteenths genes from that breed were considered to be purebred.
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Calves were typically born over an average period of 101 d between mid February and early June. Only single-birth and spring born calves were considered. Preweaning management, pasture description, and supplemental feeding of cows and calves have been previously described by Gregory et al. (1991) and Snowder et al. (2005).
Health Management and Pen Environment
Feeding and management in the feedlot were described by Gregory et al. (1994a, b). Immediately after weaning, calves were placed in the feedlot pens. Animals were begun on a diet of 2.65 Mcal of ME/kg of dry matter with 15.4% crude protein. The finishing diet consisted of 2.82 or 3.07 Mcal of ME/kg of dry matter with 11.5% crude protein. On average across years, animals were fed the starter or backgrounding diet for approximately 5 wk to become adjusted to the finishing diet. The average feeding period was 200 d, with an ADG of 1.0 kg for all steers and heifers (Table 2). Breed-type effects on growth and efficiency of gain have been previously reported (Gregory et al., 1994a,b, 1995; Rios-Utrera et al., 2005).
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). Average ages at weaning of Braunvieh, Pinzgauer, Red Poll, Simmental, and Limousin calves were less than those of the other breed types because the research protocol set a weaning age of approximately 150 d. A drought in 1988 caused calves to be weaned at an average age of 127 d. Average overall weight on feed was 205 kg but varied by breed. The average calendar day on feed was September 29. The ratio of heifers to steers in the feedlot was 49.5:50.5. Calves were fed separately by sex in mixed breed-type groups. Steers were stratified to pens according to BW. Pens were of 2 sizes, 15 x 61 m (915 m2) and 29 x 61 m (1,769 m2). Periodic removal of steers for serial slaughter caused the number of calves per pen to range from 6 to 72. There was an average of 17 steers/pen. Heifers were generally fed in the larger pens with more animals per pen in multiple breed-type groups.
All pens were in close proximity at a central feedlot location. The feedlot consisted of 2 rows of 34 pens divided by a roadway. Rows ran in an east-west direction. Additional animals not related to this study were fed in nearby pens. Feed bunks were not shared among pens, but water troughs were shared among adjacent pens. Fence line contact was possible on 2 sides of each pen, except for the end pens.
Calves were often moved among pens for consolidation and veterinary care. At approximately 90 d on feed or halfway through the feeding period, 74% of all calves were still in their initial pens. On average, 43% of all calves remained in their initial pens over the entire feeding period. Therefore, pen assignments could not be used as a classification effect.
Calves were vaccinated before weaning and at weaning before entry into the feedlot. The vaccination program varied across years. Killed virus vaccines used from 1987 to 1992 were replaced with modified live virus vaccines in 1993. A parainfluenza-3 vaccine was added to the calf vaccination program in 1994. At approximately 42 d of age, calves were vaccinated with an 8-way Clostridial and a 5-way Leptospirial vaccine. At approximately 165-d of age, calves were given booster vaccinations of Clostridial and Leptospirial vaccines and a modified live virus vaccine for infectious bovine rhinotracheitis, parainfluenza-3, and bovine viral diarrhea.
It is not known whether the epidemiology of BRD observed in this study resembles that of commercial feedlots. Numbers of calves/pen and movement of calves among pens in commercial feedlot enterprises would differ from those in this study. Commercial feedlots also receive cattle from many different locations at different times, increasing the probability of introducing BRD carriers or exposing cattle to different stress factors from those with the closed animal system at US Meat Animal Research Center.
Disease Detection
Calves were monitored daily by the staff veterinarian or the beef cattle research technicians, or both. Diseases were detected by physical examination, necropsy, or laboratory analyses and were recorded. Typical clinical BRD symptoms for calves included 1 or more of the following: fever, rapid breathing, repetitive coughing, nasal or eye discharge or both, diarrhea, dehydration, and appetite depression. Disease codes related to BRD were combined. The majority of the BRD classifications were from codes for pneumonia and respiratory disease (86 and 11%, respectively). Additional disease codes included for BRD were bronchitis, emphysema, pleuritis, pulmonary adenomatosis, upper respiratory infection, and pleural fibrosis. Because such diseases may be interpreted as BRD by feedlot staff, it seemed appropriate to include them with the BRD codes. The recoded binary classification listed calves as healthy (code = 200) or affected by BRD (code = 100). Records also included calves that died due to BRD symptoms. To avoid multiple measurements on the same calf, which may have been due to lingering respiratory disease or reinfection, only the initially observed infection during the feedlot period was considered.
Additional vaccinations were administered to calves diagnosed with BRD or other ailments. For calves expressing clinical signs of BRD, the calf’s age, weight, severity of clinical symptoms, and drug withdrawal time were considered in determining the appropriate treatment. All treatments for BRD included administration of 1 or more medications (oxytetracycline, ceftiofur, flunixin meglumine, florfenicol, tylosin, enrofloxacin, and sulfadimethoxine). Calves with BRD were checked daily and may have received several therapy treatments, as determined by the veterinarian or technical staff, or both. Antibiotics were not routinely administered in the calves’ feed or water.
Approximately 13% of all calves were diagnosed with BRD during the preweaning period. Because it is not known if BRD infection during the preweaning period affects BRD incidence in the feedlot, a binomial trait for previous exposure was created.
Classification could not be made as to whether an animal was tolerant or resistant to BRD in the feedlot. Tolerance is defined as the ability of an infected animal to show little or no measurable detrimental effect of the disease. Resistance is the ability of the animal to resist infection or control the lifecycle of the infecting microbe. In this study, "resistant" was used to describe an animal that was not detected with clinical symptoms of BRD without regard to whether the animal was truly tolerant, resistant, or not exposed to BRD-causing organisms. The actual trait measured was treatment of observed clinical BRD.
Statistical Procedures
The pedigree used to calculate the relationship matrix consisted of 47,880 animals born between 1972 and 2001, including 1,655 sires and 15,648 dams. The number of animals in the pedigree with a nonzero inbreeding coefficient was 6,984. The average inbreeding coefficient was 2.3%, and ranged from 0.00 to 0.25 (only 161 animals with an inbreeding coefficient greater than 0.10). Total numbers of animals in the pedigree by breed type are reported in Table 3. Of the 18,112 feedlot calves, 5,104 had a nonzero inbreeding coefficient, with an average of 2.2%. The effect of such small levels of inbreeding on BRD incidence was assumed to be negligible.
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and 5. Within-breed-type statistics for the percentage of animals with a preweaning BRD record, average age and days on feed when BRD was observed, incidence, mortality, and total death loss (mortality of BRD calves expressed as a percent of all calves) are shown in Table 5.
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Variance Components
Data from all purebred and composite breeds were combined into an overall analysis of BRD as the dependent variable. Because some breed types were associated with years, and incidence of BRD was greater from 1987 to 1992 (Figure 1), the data were also partitioned into 3 subsets (Table 1). The first set (EARLY) consisted of all 12 breed types from 1987 to 1991 (n = 7,039). The second set (LATE) included the 7 breed types present from 1992 to 2001 (n = 9,981) when the overall incidence of BRD was less than in previous years. The Hereford breed, although not present in 2000 and 2001, was included in this latter data set. The third set (SELECT) included only the 6 breed types present in all years from 1987 to 2001, plus Herefords (n = 16,208).
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), analyses of EARLY and LATE subsets were performed, respectively, within periods of high (1987 to 1991) and low (1993 to 2001 to 1992 exception) BRD incidence. Analysis of the SELECT subset permits appropriate individual breed-type contrasts among the 4 purebred and 3 composite breed types across all years. This latter analysis may be more informative of breed-type differences than analysis of the LATE subset. The effect of heterozygosity was inferred by contrasting individual composite breeds with each pure breed. More appropriately, composite breeds were further contrasted with all purebreds using appropriate coefficients for the purebreds based on their total contribution to the composite breeds.
Variance components for incidence of BRD were estimated using a single-trait model with a derivative-free REML algorithm (Graser et al., 1987) by using the computer programs of Boldman et al. (1995). Convergence was considered to have been reached when the variance of the –2 logL in the simplex was less than 1 x 10– 6. After initial convergence, 4 restarts were performed to ensure global convergence as determined when the –2 logL did not change to the second decimal. Standard error of the heritability estimate was based on the average information matrix and the delta method (e.g., Dodenhoff et al., 1998). Fixed effects in the model included year, breed type, and sex of calf. Initial weight on entering the feedlot and total days on feed were included as covariates. Calf (genetic direct) was a random effect. Because breed types were confounded with years, and incidence of BRD varied with groups of years, breed type by year was regarded as an uncorrelated random effect. This approach allows the SE to account for the interaction variance and the residual variance, which in turn requires a much larger difference in breed-type solutions to be significant. All subsets were analyzed with the same model. Contrasts between solutions for sex of calf and breed types for BRD incidence were tested with a Student’s t-test. Individual breed-type solutions were deviated from the solution for the Angus breed.
Because BRD incidence is a dichotomous threshold trait and because not all animals were likely to have had equal opportunity to become infected and express their true phenotype, it is reasonable to assume there is an underlying scale of genetic variation (Robertson and Lerner, 1949). The overall estimate of heritability for BRD on the observed scale was transformed to a heritability estimate on the underlying continuous normal scale (Robertson’s appendix to Dempster and Lerner, 1950).
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RESULTS AND DISCUSSION |
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TopAbstractINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONIMPLICATIONSLITERATURE CITED |
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).
Characteristics of BRD
The incidence of BRD varied across the 15-yr span (Figure 1) and ranged from 4.6 to 43.8% per year with an average annual incidence of 17.0%. The greatest incidences were from 1987 to 1992. Epidemic proportions were observed in 1990 and 1991. In 1990, observed incidences within breed type ranged from 28 to 73% (Braunvieh and Hereford, respectively; Table 4). Several factors may have contributed to greater incidences in earlier years. These factors included the use of less effective killed vaccines from 1987 to 1992 compared with modified live vaccines used from 1993 to 2001, adding parainfluenza-3 vaccine to the calf vaccination program in 1995, and changing the castration method from surgical (1987 to 1989) to mechanical banding (1990 to 2001). After 1992, the annual overall incidence remained less than 14%, although there were occasional high incidences for some breed types such as 20 and 21% for Charolais and MARC I in 1994 and 26% for Gelbvieh in 1996 (Table 4). The average incidence of BRD in 1999 for 520 US commercial feedlots was reported to be 14.4% (NAHMS, 2000a) and to be 20.6% for 17 Iowa feedlots from 1988 to 1997 (Faber et al., 1999). Large variations in morbidity due to BRD ranging from 0 to 59% were reported in Iowa for 20 feedlots across 10 yr (Faber et al., 1999).
The average age when feedlot calves were treated for BRD was 202 ± 51 d and ranged from 109 to 522 d. The number of calves with BRD treated per day across all years was associated with days on feed (Figure 2). A dramatic increase in incidence was observed after 5 d in the feedlot. Incidence of BRD peaked within 14 d on feed and remained high until approximately 80 d on feed. After 110 d on feed the numbers of calves observed with BRD per day were negligible. The epidemiological pattern observed is similar to that previously reported in feedlot cattle by Loneragan (2001) for BRD and by Schunicht et al. (2003) for undifferentiated disease treatment.
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). Effects of breed type and environment on incidence of BRD in calves before weaning have been previously reported (Snowder et al., 2005). Preweaning incidence was greatest in Braunvieh calves and lowest in Hereford calves. Among feedlot calves with previously observed BRD, 18% (n = 417) were also detected with BRD in the feedlot. Among feedlot calves with no previous record of BRD (n = 15,794), 16% (n = 2,524) were detected with BRD. The effect of previous BRD infection was not a significant effect when analyzing feedlot traits for ADG and BRD incidence (P = 0.36 and 0.24, respectively).
Effect of Sex of Calf
Overall incidence of BRD differed (P < 0.05) between steers and heifers (20 vs. 14%, respectfully). During years of high incidence (1987 to 1991), male calves made up approximately 62% of all reported cases of BRD but only 54% during years of low incidence (1992 to 2001). Castration before entry into the feedlot may predispose male calves to BRD infection. The change in castration procedure in 1990 may partially account for the difference in BRD incidence between the 2 periods. From an extensive review of literature, Bretschneider (2005) concluded that age of castration had a significant effect on ADG for the first 30 d postcastration and on stress response, and that rubber banding methods were less stressful than surgical castration. Management of steers differed slightly with fewer animals per pen and more frequent weighing than heifers. The interaction of breed type and sex of calf effects was not significant in preliminary analyses.
Effect of Breed Type
Unadjusted averages for traits related to BRD varied among breed types (Table 5). Average age when BRD was detected did not vary greatly among breed types with an overall average of 202 d. Limousin and Simmental had the youngest average ages (190 d), whereas Charolais had the oldest average age (213 d). The average number of days on feed when BRD was detected was 43 d, ranging by breed type from 35 to 56 d (Angus and Braunvieh, respectively).
The unadjusted mean incidence for BRD was lowest for Angus, 10.2% (Table 5). Charolais, Gelbvieh, MARC I and III cattle also had lower unadjusted mean incidences. Incidence of BRD was greatest for Pinzgauer, Braunvieh, Simmental, and Limousin (35, 34, 33, and 32%, respectively).
When only years of 1987 through 1991 (EARLY) were considered (the year for which records were available for all 9 pure breeds and 3 composites), high incidences were observed in all breed types. Incidence by year and breed type ranged from 7 to 73% (Table 4). Solutions to the mixed model equations indicated that the only significant differences were between MARC I calves and Hereford and Pinzgauer calves (Table 6).
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). When breed types present in all years (1987 to 2001, SELECT) were considered, the large solution for Herefords differed (P < 0.05) from solutions for Angus, MARC I, and MARC II (Table 8). The extremely large incidence of 73% in 1990 (Table 4) for the Herefords may be a main reason for these differences. Greater incidences of BRD in the Herefords during 1998 and 1999 (19 and 15%, respectively) compared with lower incidences in Angus, MARC I, and MARC II calves (2 to 7%; Table 4) may have influenced these differences.
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). Approximately 4% of calves detected with BRD died, which contributed to a total death loss of 1%. In a national feedlot monitoring program, the proportion of total death losses attributed to BRD increased from 52.1% in 1994 to 61.5% in 1999 with an average annual total death loss of 1.2% (NAHMS, 2000b). Changes in the cattle and feedlot industries may have contributed to the increased death loss observed nationally such as younger calves being fed longer, lack of knowledgeable and experienced feedlot labor to detect BRD, lack of preweaning vaccinations, and negligible prevention at the producer level (Ishmael, 2001).
Mortality of calves detected with BRD was greatest in the Red Poll (8.9%). Mortality of Red Poll calves before weaning attributed to BRD was previously reported to be greater than for most other breed types (Snowder et al., 2005). The greater mortality rate among Red Poll calves before weaning and during the feedlot period suggests this breed may be more sensitive to clinical BRD infection compared with other breed types.
No apparent advantage for heterozygosity to BRD resistance (incidence or mortality) could be detected from individual contrasts of composite breeds with each pure breed. Although the solutions for MARC I and III composite calves were favorable for resistance compared with most other breed types (Table 6, 7, and 8), those composite breeds were not less susceptible than other breed types (P > 0.05) except when compared with the Herefords. The group contrast of all 3 composites with all 9 pure breeds was not significant (P > 0.05). Therefore, heterozygosity cannot be considered as a significant factor on resistance to BRD.
Heritability of BRD
Estimates of heritability of resistance to BRD in the feedlot were low, ranging from 0.04 ± 0.01 to 0.08 ± 0.01 (Table 9). No previous estimates could be found in the literature. However, these estimates are similar to estimates reported for calves before weaning by Muggli-Cockett et al. (1992, 0.10 ± 0.02) and Snowder et al. (2005, 0.07 ± 0.01 to 0.19 ± 0.01). The larger estimate for the EARLY data set (years of high incidence) compared with the smaller estimate from the LATE data set (years of low incidence) supports the conclusion by Snowder et al. (2005) that greater heritability estimates are related to greater incidences as more animals are challenged to express their phenotype for disease resistance.
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), and the overall annual average BRD incidence of 17% (Table 5). The larger heritability estimate for the underlying continuous scale implies selection for BRD could be effective provided all animals are challenged adequately. Thus, development of an inexpensive and humane method of challenging animals for BRD resistance would seem an important step in developing a selection program to reduce incidence of BRD.
The results of this study suggest that selection against susceptibility to BRD using producer or industry recorded data is feasible, which agrees with conclusions of previous reports that producer-recorded data can be useful for genetic selection to reduce diseases, reproductive disorders, and metabolic disorders (Zwald et al., 2004a,b).
Implications to Management
Although BRD is a disease of foremost concern because of its widespread incidence in cattle (Callan and Garry, 2002), BRD can be controlled by decreasing pathogen transmission between animals. Management practices that reduce pathogen introduction, exposure, and transmission are important initial steps. Total eradication of pathogens for BRD is not likely in feedlot environments where introduction of cattle and movement of cattle are usual management activities. Decreasing pathogen transmission between animals can be achieved by removing affected animals and avoiding crowding of animals. Another approach is to reduce the number of susceptible animals by increasing the number of BRD-resistant animals, which can be achieved by the use of vaccines or antibiotics, although some pathogens may be resistant to antibiotics and efficacy of vaccines may be limited in some cases. Vaccines can be effective for reducing not only susceptibility but also for reducing shedding of infectious BRD agents to other calves (Frank et al., 1994, 2003). In some cases antibiotics will not make the animal resistant but, for example, can reduce carriage and shedding of M. haemolytica (Frank et al., 2000, 2002). Another approach is to select for individuals resistant to BRD. All animals in a population need not be resistant to prevent an epidemic if enough animals resistant to the disease are present to prevent spread to susceptible animals (Anderson and May, 1992; Bishop and MacKenzie, 2003).
Pathogens resulting in BRD in this study were probably not of a single variety, but rather could have been of bacterial, viral, or mycoplasmal origin, or some combination of these. If this assumption is correct, then selection for BRD resistance or tolerance based on common treatment or incidence records may result in an improved generalized disease defense mechanism, possibly a more efficacious immune system. Previous studies demonstrated that selection for antibody or cell-mediated immune responses, or both, improve health and production in cattle (Hernández et al., 2003), swine (Mallard et al., 1992 and 1998), and poultry (Pinard-van der Laan et al., 1999). Further investigation of the genetic mechanisms of disease resistance is certainly warranted.
Economic Effect
The general economic effect of BRD on feedlot cattle was determined by its effect on ADG and calf death loss, ignoring treatment costs. All fixed factors in the linear model to examine the effect of BRD on ADG were significant with the model accounting for 80% of the variation. Calves diagnosed with BRD had lower (P < 0.001) ADG (0.95 kg) compared with healthy animals (0.99 kg). Over a 200-d feeding period, an 8-kg difference between a healthy and an infected calf could be expected. This is in agreement with Bateman et al. (1990) who observed a 0.06 kg difference for ADG between healthy and calves treated for BRD on feed for 138 to 235 d. However, Faber et al. (1999) reported that after cattle were effectively treated for BRD, compensatory gain was sufficient such that no difference in ADG was observed over the entire 220-d feeding period. Compensatory gain by BRD-affected calves was also inferred by Bateman et al. (1990) who reported that ADG of BRD affected calves was 0.14 kg less than healthy calves after 28 d but that this difference diminished to 0.06 kg after 135 to 238 d on feed. Because the greatest incidence of BRD generally occurs early in the feedlot, prompt detection and proper treatment would contribute to a calf’s recovery and subsequent compensatory gain.
A feedlot with 1,000 calves and a 14.4% incidence (NAHMS, 2000a) could expect a loss of 1,152 kg plus treatment costs attributed to BRD. Based on the current slaughter cattle price of $1.83/kg (USDA, AMS, 2005), the value in lost gain attributed to BRD would be $2,108 per 1,000 animals, assuming calves are slaughtered after 200 d on feed. Additionally, if treatment costs for medication and related items averaged $15.57 per sick animal (Faber et al., 1999; USDA APHIS, 2001), then an additional loss of $1,813 could be attributed to treating 144 infected calves. A death loss of 1% due to BRD (Table 5) would cost $9, 974 (10 calves with average market weight of 545 kg at slaughter at $1.83 kg, USDA AMS, 2005). Estimated total economic loss per 1,000 animals would be $13,895, not including feed costs before the death of calves, labor and associated handling costs. A survey of cattle feedlots in Iowa estimated that steers not infected with BRD returned $57.48 more per steer (Faber et al., 1999). Overall economic loss due to BRD may be difficult to determine because animals not exhibiting overt clinical symptoms can have suppressed gains (Wittum et al., 1996).
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IMPLICATIONS |
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TopAbstractINTRODUCTIONMATERIALS AND METHODSRESULTS AND DISCUSSIONIMPLICATIONSLITERATURE CITED |
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1 Corresponding author: snowder{at}email.marc.usda.gov
Received for publication January 25, 2006. Accepted for publication February 27, 2006.
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LITERATURE CITED |
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