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J. Anim Sci. 2008. 86:2082-2088. doi:10.2527/jas.2007-0523
© 2008 American Society of Animal Science

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ANIMAL GENETICS

Genetic components of heat stress in finishing pigs: Development of a heat load function

B. Zumbach*,{dagger},1, I. Misztal*, S. Tsuruta*, J. P. Sanchez*,{ddagger}, M. Azain*, W. Herring§, J. Holl§, T. Long§ and M. Culbertson§

* Department of Animal and Dairy Science, University of Georgia, Athens 30602-2771; and {dagger} Norsvin, Pb 504, 2304 Hamar, Norway; and {ddagger} Departamento de Producción Animal, Universidad de León, León, 24071, Spain; and § Smithfield Premium Genetics Group, PO Box 668, Rose Hill, NC 28458


    Abstract
 Top
 Abstract
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 LITERATURE CITED
 
The objective of this study was to quantify the effect of heat stress during the life of a pig on its final weight, as a first step toward a genetic evaluation for heat tolerance. Data included carcass weights of 23,556 crossbred pigs [Duroc x (Landrace x Large White)] raised on 2 farms in North Carolina and slaughtered from May 2005 through December 2006. Weather data were available from a nearby weather station. Lifetime of a pig was assumed to be partitioned into 2 periods. During an initial period, the effect of heat stress was assumed to be negligible or compensated for later. During the second period ending in slaughtering, the ADG was assumed to be affected linearly by heat load. Weekly heat load was calculated as degrees of average temperature-humidity index in excess of a threshold (18°C). The total heat load (H) was the sum of weekly heat loads during the second period. During the months of January to May H was 0; H reached a peak in September. The final BW during the peak of heat stress decreased about 6 kg compared with BW during months of non-heat stress. Weekly and monthly averages of carcass weight generally moved similarly to H. However, there were large fluctuations unrelated to H; the fluctuations were different on the 2 farms. The model included the effects of farm-year of slaughter, sex, age at slaughter, and H, where age at slaughter and H were linear regressions. In analyses, the threshold was varied from 16 to 20°C, and the second period was varied from 8 to 16 wk. The greatest R2 (10.4%) was at the threshold of temperature-humidity index = 18°C for a period of 10 wk. Varying the threshold and the length of time reduced R2 less than 1%. Least squares means of year-month and year-week of carcass weight were calculated using a model with the fixed effects farm-year-month or farm-year-week of slaughter, sex, and age at slaughter (linear covariate), and the random effect of birth litter. Changes in BW of finisher pigs due to heat stress can be quantified by H during the last 10 wk of the life of the pig.

Key Words: carcass weight • heat stress • pig


    INTRODUCTION
 Top
 Abstract
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 LITERATURE CITED
 
Heat stress in pigs impairs not only the economics of the pig industry (e.g., St-Pierre et al., 2003Go) but also the animals’ welfare and environment (Huynh, 2005Go). Due to selection for leanness, total heat production of growing pigs has increased during the last decades (Brown-Brandl et al., 2001Go). The more heat an animal produces internally by its metabolism, the smaller its capacity for tolerating external heat (Bianca, 1976Go).

Response to heat stress begins with increased respiration rate, continues with decreased feed intake, and leads to increased rectal temperature (Huynh, 2005Go). Although reduced feed intake can be considered a mechanism of thermoregulation (Bianca, 1976Go), increased rectal temperature is an indicator of exhaustion of the thermoregulation capacity of pigs (Huynh, 2005Go). In general, if the magnitude (intensity and duration) of potential stressors exceeds a threshold, animals are unable to cope and are affected adversely (Hahn et al., 2003Go).

For production in a hot environment, especially in a commercial environment where management conditions are not as good as on nucleus farms, it would be desirable to select pigs resistant to heat stress. Such selection could be for the lowest rectal temperatures during the hot season. However, such data are not readily available. Ravagnolo et al. (2000)Go used weather records from public weather stations to quantify the change in milk yield in response to the changing temperature-humidity index (THI). They also used the same records to determine genetics of heat tolerance (Ravagnolo and Misztal, 2000Go). Records from nearby public weather stations were found to be as accurate in capturing the effect of heat stress as records collected on farm (Freitas et al., 2006Go). As opposed to dairy cattle, where heat stress on 1 d affects milk yield 1 or 2 d later (Fuquay, 1981Go; Ravagnolo et al., 2000Go), the effect of daily heat stress on growth is more difficult to measure. The purpose of this study was to quantify the effect of heat stress during the life of a pig on its final BW, as a first step toward a genetic evaluation for heat tolerance.


    MATERIALS AND METHODS
 Top
 Abstract
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 LITERATURE CITED
 
Animal Care and Use Committee approval was not obtained for this study because the data were from an existing database.

Data

Records on warm carcass weight data were available on 23,556 terminal crossbred pigs [Duroc1 x (Landrace x Large White)] slaughtered from May 2005 through December 2006 (Smithfield Premium Genetics Group, Rose Hill, NC). Duroc1 corresponds to the Duroc nucleus line P1 (Zumbach et al., 2007Go). The animals were raised on 2 commercial, 1,200-sow farrow-to-finish farms.

Farm 1 provided 12,771 records, whereas farm 2 contributed 10,785 records. Both farms were located in North Carolina about 8 km apart. The average age at slaughter was 200 d, and the average carcass weight was 89 kg. There was no evidence of a seasonal pattern for age at slaughter (Table 1Go). Typical and similar corn-soybean diets were used at both sites. Feeding was ad libitum for all animals.


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Table 1. Number of observations, means (SD) of age at slaughter, and carcass weight per month of slaughter for each farm
 
Because no inside temperature-humidity data were available, these were obtained from a public weather station located at an airport within a distance of 21 km and 14 km from farm 1 and farm 2, respectively. Average daily temperature (T) in degrees Celsius (°C) and relative humidity (RH) based on hourly observations were available (State Climate Office of North Carolina, 2007Go). A THI (NOAA, 1976Go) for each day (average of 24-h period) of the experimental period was calculated based on the formula:


Formula

Because a single day of heat stress could be compensated by following cooler days, THI was calculated on a 7-d basis (rolling average) as the average of daily THI.

Quantification of Heat Load

Let heat load in a unit of time (h) be:


Formula

where THI = average THI for a unit of time and THI0 = threshold of heat stress (upper critical temperature). Let tij be THI of animal i at age j. Assume that any effect of heat stress up to age p is compensated later and that growth after age p is constant, except being linearly affected by heat load. Then, the weight at time of slaughter s is:


Formula

where weightis = BW of animal i at slaughter and adgi = ADG of animal i. This can be simplified to:


Formula

where weighti0 = slaughter weight of animal i in the absence of heat load and Hi = total of heat load for that animal in the time interval from p+1 to s.

Daily heat load was defined as the number of degrees exceeding the threshold. The weekly heat load was defined as a mean of daily THI exceeding the threshold. The total heat load (H) was defined as the sum of weekly heat loads over the last n weeks of life, where n is a constant.

Both the threshold and the number n were selected based on the greatest coefficient of determination (R2) obtained with the following model:


Formula

where yijklm = carcass weight on farm-year i, sex j, with slaughter age k standardized to 200 d, and heat load l. The fixed effects were FYi = farm-year, i = 1 to 4; Sj = sex, j = 1, 2; b1 = regression coefficient 1; agek = age at slaughter of the kth animal, k = 1 to 21,653; b2 = regression coefficient 2; Hl = heat load at slaughter date, l = 1 to 103; eijklm = the residual. This model could not include the day of slaughter because of confounding with heat load.

Seasonal Effects

Least squares means for year-month of slaughter and year-week of slaughter were calculated according to the model:


Formula

where yijklm = carcass weight on farm-year-week or farm-year-month of slaughter date i, sex j, with slaughter age k standardized to 200 d, and birth litter l. The fixed effects were FYWi/FYMi = farm-year-week of slaughter date, i = 1 to 170/farm-year-month of slaughter date, i = 1 to 40; Sj = sex, j = 1, 2; b = regression coefficient; agek = age at slaughter of the kth animal, k = 1 to 21,653. The effects ll = birth litter, l = 1 to 4,305, and eijklm = residual were random. The number of observations per slaughter month and farm are shown in Table 1Go.


    RESULTS AND DISCUSSION
 Top
 Abstract
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 LITERATURE CITED
 
The coefficients of determination (R2) considering duration of heat stress from 8 to 16 wk before slaughter and different THI thresholds are presented in Figure 1Go. The R2 ranged between 9.55 and 10.45% showing maxima for the period of 10 wk for all THI thresholds considered. The decrease to the right (more weeks considered) was slower than to the left (shorter period), and the optimum was broad (e.g., considering 9 to 13 wk of heat stress, R2 varied only about 0.2%). The curve with THI0 = 18°C was on the greatest level followed by THI0 = 16°C. At the maximum, R2 was similar for the different combinations. It differed less than 0.1% between THI0 = 18 and 16°C and about 0.2% between THI0 = 18 and 20°C.


Figure 1
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Figure 1. The R2 for different heat load functions depending on critical temperature-humidity index (THI0) and time period considered previous to slaughter (no. of weeks).

 
The effect of THI0 on H, considering a heat stress period of 10 wk, is shown in Figure 2Go. The maxima of H were at the beginning of September in both years. They coincided for the 3 THI0 considered. The period of non-heat load (H = 0) was from January to April-May 2006. The lower the THI0, the earlier was the onset of H. For example, the onset of H in 2005 for THI0 = 20°C was in the beginning of June. At that time, H had reached 6 and 14% of the maximum, for THI0 = 18°C and THI0 = 16°C, respectively. The end of the heat stress period (H = 0) was at about the same time for all of the thresholds considered.


Figure 2
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Figure 2. Heat load with different critical temperature-humidity indexes (THI0) considering a period of 10 wk (#wk) previous to slaughter.

 
The heat load curves at THI0 = 18°C for different periods are presented in Figure 3Go. The onset of H was identical for the periods of 8, 10, and 12 wk. The maxima of the 2-wk longer periods were delayed by about 1 wk, whereas the offsets were prolonged by about 2 wk. In experimental studies with controlled heat regulation inside the stable, the upper critical temperature for voluntary feed intake in pigs was between 22.9 and 25.5°C (Huynh et al., 2005Go); Quiniou et al. (2001)Go and Le Bellego et al. (2002)Go considered 23 and 24°C, respectively, as thermoneutrality.


Figure 3
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Figure 3. Heat load considering different periods (#wk = number of weeks) previous to slaughter at a critical temperature-humidity index (THI0) of 18.

 
While THI from the weather station accounted for the outside climate, the pigs were kept within partially open barns. This would explain the relatively low THI0. Based on the R2 evaluation, the THI0 for the fixed heat load model was set to 18°C, and the optimum number of weeks considered before slaughter was 10.

The optimum THI0 was 18°C and was lower than thresholds found elsewhere. This could have been due to the definition of THI as a 24-h average, which was additionally averaged over a period of 1 wk. A 24-h average would be greater than a 2-d average during the peak of heat stress, when THI drops very slowly, and lower at the beginning or the end of summer, when nightly cooling occurs more quickly. Studies to identify the most suitable THI in dairy cattle used THI based on maximum temperature and minimum humidity and the average temperature and humidity (Ravagnolo et al., 2000Go) or THI with different weights for temperature and humidity for assessing losses in milk performance (Bohmanova et al., 2007Go).

The least squares means of carcass weight across the months of slaughter and the computed heat load for the 2 farms are presented in Figure 4Go. Weights for farm 1 had large fluctuations that appeared to be non-heat stress related. On both farms, drops in carcass weight occurred during August through November in 2005 and July through October in 2006. The difference in carcass weight between heat stress and non-heat stress months was about 6 kg. The 2 farms had an identical design and were located about 8 km apart. Differences between the farms were due to management and random factors that could not be accounted for in this study.


Figure 4
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Figure 4. Least squares means of year-month of slaughter for carcass weight of pigs raised on farms 1 and 2 from May 2005 through December 2006.

 
Figure 5Go presents the weights using combined data from the 2 farms and the H curve. The months with the lower carcass weights coincided with those of increased heat load. The onset and recovery from heat stress in 2005 seemed to occur 1 mo later than the increase and decrease of H, whereas in 2006, H and carcass weight were in agreement (see also Figure 6Go). In Figure 6Go, the carcass weights and H are shown on a weekly basis. Although the fluctuations between weeks of slaughter were large, the pattern of the curve coincided with the H curve. In Figure 7Go, a scatter plot of weekly carcass weights against H shows the tendency of decreasing carcass weights with increasing H.


Figure 5
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Figure 5. Least squares means of year-month of slaughter for carcass weight (CW) and heat load (H) from May 2005 through December 2006.

 

Figure 6
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Figure 6. Least squares means of year-week of slaughter for carcass weight (CW) and heat load (H) from May 2005 through December 2006.

 

Figure 7
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Figure 7. Scatter plot of least squares means of year-week of slaughter for slaughter weight against heat load during the study period.

 
The effect of heat load on carcass weight was due to many factors, including heat management, as well as sensitivity to heat stress at preweaning and postweaning, compensatory gain after a decline in gain due to heat stress, and both short- and long-term effects of heat stress. Our definition of H involved many assumptions including no uncompensated detrimental effect on growth due to heat stress during the first 4.5 mo of life (i.e., until 10 wk before slaughter). It also assumed linear growth for the finishing pigs during non-heat stress seasons, which was in agreement with the results of Knap et al. (2003)Go and Edwards et al. (2006)Go.

Because humidity combined with high temperature affects pig growth (Huynh et al., 2005Go), THI seemed to be an appropriate variable for the definition of H. With the summation of THI above thermoneutrality, our definition of H accounted for both severity and duration of heat stress. The model did not allow for compensatory growth for heat stress occurring up to 10 wk before slaughter. This seemed to hold, because pigs were kept under commercial conditions where several stressors such as stocking rate or disease pressure are acting simultaneously. Kerr et al. (2005)Go reported greater live weight gain after the heat stress application for pigs housed individually at low density compared with pigs maintained at high density. The effect of disease challenge was reported to be more profound than that of heat stress (Kerr et al., 2003Go). The fluctuations in the carcass weight curves in this study indicated the existence of stressors different from heat. These occurred without any seasonal pattern throughout the study period. No seasonal pattern could be detected for mortality (unpublished data). Although the function of heat load provided only a limited fit to real carcass weights, the limitation was due to lack of data. A study of more accurate accounting for heat stress will be undertaken when more data with smaller weekly-monthly fluctuations become available.

In conclusion, reduced carcass weight in pigs due to heat stress can be quantified by the combined heat load during 2 to 3 mo preceding slaughter. In North Carolina, these reductions were present in weights of pigs slaughtered from July to November. The quantification can be used for determining the quality of management, properties of particular lines with respect to heat stress, and possibly in genetic selection toward heat tolerance.

1 Corresponding author: Birgit.zumbach{at}norsvin.no

Received for publication August 16, 2007. Accepted for publication April 21, 2008.


    LITERATURE CITED
 Top
 Abstract
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 LITERATURE CITED
 


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Bohmanova, J., I. Misztal, and J. B. Cole. 2007. Temperature-humidity indices as indicators of milk production losses due to heat stress. J. Dairy Sci. 90:1947–1956.[Abstract/Free Full Text]

Brown-Brandl, T. M., R. A. Eigenberg, J. A. Nienaber, and S. D. Kachman. 2001. Thermoregulatory profile of a newer genetic line of pigs. Livest. Prod. Sci. 71:253–260.[CrossRef]

Edwards, D. B., R. J. Tempelman, and R. O. Bates. 2006. Evaluation of Duroc- vs. Pietrain-sired pigs for growth and composition. J. Anim. Sci. 84:266–275.[Abstract/Free Full Text]

Freitas, M. S., I. Misztal, J. Bohmanova, and J. West. 2006. Utility of on- and off-farm weather records for studies in genetics of heat tolerance. Livest. Sci. 105:223–228.[CrossRef]

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Huynh, T. T. T. 2005. Heat stress in growing pigs. PhD Thesis. Wageningen Univ., the Netherlands.

Huynh, T. T. T., A. J. A. Aarnink, M. W. A. Verstegen, W. J. J. Gerrits, M. J. W. Heetkamp, B. Kemps, and T. T. Canh. 2005. Effects of increasing temperatures on physiological changes in pigs at different relative humidities. J. Anim. Sci. 83:1385–1396.[Abstract/Free Full Text]

Kerr, C. A., G. J. Eamens, J. Briegel, P. A. Sheehy, L. R. Giles, and M. R. Jones. 2003. Effects of combined Actinobacillus pleuropneumoniae challenge and change in environmental temperature on production, plasma insulin-like growth factor I (IGF-I), and cortisol parameters in growing pigs. Aust. J. Agric. Res. 54:1057–1064.[CrossRef]

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Quiniou, N., J. Noblet, J. van Milgen, and S. Dubois. 2001. Modelling heat production and energy balance in group-housed growing pigs exposed to low and high ambient temperatures. Br. J. Nutr. 85:97–106.[Medline]

Ravagnolo, O., and I. Misztal. 2000. Genetic component of heat stress in dairy cattle, parameter estimation. J. Dairy Sci. 83:2126–2130.[Abstract]

Ravagnolo, O., I. Misztal, and G. Hoogenboom. 2000. Genetic component of heat stress in dairy cattle, development of heat index function. J. Dairy Sci. 83:2120–2125.[Abstract]

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B. Zumbach, I. Misztal, S. Tsuruta, J. P. Sanchez, M. Azain, W. Herring, J. Holl, T. Long, and M. Culbertson
Genetic components of heat stress in finishing pigs: Parameter estimation
J Anim Sci, September 1, 2008; 86(9): 2076 - 2081.
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