You are on page 1of 9

Livestock Production Science 63 (2000) 245253 www.elsevier.

com / locate / livprodsci

Voluntary feed intake and feeding behaviour of group-housed growing pigs are affected by ambient temperature and body weight q
1 N. Quiniou , S. Dubois, J. Noblet*

Station de Recherches Porcines, Institut National de la Recherche Agronomique, 35590 Saint-Gilles, France Received 25 September 1998; received in revised form 7 April 1999; accepted 21 June 1999

Abstract The effect of ambient temperature on individual feeding behaviour was studied in six groups of Pietrain 3 Large White barrows. In experiment 1 (two groups), ambient temperature varied in a cyclic way from 22 to 128C and 12 to 228C with three or four consecutive days at each of the following temperatures: 22, 19, 16, 14 or 128C. Similarly, in experiment 2 (two groups), temperature varied from 19 to 298C and 29 to 198C with three or four consecutive days at 19, 22, 25, 27 or 298C. In both experiments, each group was used over two successive cycles with an initial body weight (BW) of 37 kg at cycle 1 (four pigs per group) and 63 kg at cycle 2 (three pigs per group). During experiment 3, groups of four pigs were exposed to varying temperatures over one cycle either as in experiment 1 (one group) or as in experiment 2 (one group); their initial BW was 45 kg. Photoperiod was xed to 12 h of light. In experiments 1 and 2, neither the daily number of meals (11) nor the rate of feed intake (37 g / min) were affected by temperature. The daily number of meals was lower at cycle 2 (9 vs. 12 at cycle 1 on average) but their size was higher (305 vs. 181 g / meal at cycle 1). The feeding pattern was mainly diurnal (62%). From individual data obtained at each temperature level and each stage of growth in this study (N 5 296), an equation to predict the voluntary feed intake (VFI) from temperature (T, ranging between 12 and 298C) and body weight (BW, ranging between 30 and 90 kg) is proposed: VFI (g / d) 5 2 1264 1 117T 2 2.40T 2 1 73.6BW 2 0.26BW 2 2 0.95T 3 BW (RSD 5 329). The present relationship indicates that VFI depends on temperature and body weight with a marked negative effect of high ambient temperatures in heavier pigs. 2000 Elsevier Science B.V. All rights reserved.
Keywords: Pig; Appetite; Temperature; Feeding behaviour

Preliminary results were presented as an abstract at the 32nd Congress of the International Society of Animal Ethology, 2125 July 1998, Clermont-Ferrand, France. *Corresponding author. Tel.: 133-2-9928-5049; fax: 133-29928-5080. E-mail address: noblet@st-gilles.rennes.inra.fr (J. Noblet) 1 On leave from the Institut Technique du Porc, BP 3, 35650 Le Rheu, France. Tel.: 133-2-9960-9838; fax: 133-2-9960-9355. E-mail: nathalie.quiniou@itp.asso.fr.

1. Introduction During the growing period, the voluntary feed intake (VFI) is inuenced by the physiological status (age, body weight) (Kanis and Koops, 1990) and the type of pig (breed, sex) (Fuller et al., 1995; Labroue et al., 1995). In addition, environmental factors, such as temperature, affect VFI. According to studies

0301-6226 / 00 / $ see front matter 2000 Elsevier Science B.V. All rights reserved. PII: S0301-6226( 99 )00135-9

246

N. Quiniou et al. / Livestock Production Science 63 (2000) 245 253

available in the literature, the decrease of temperature below the thermoneutral zone is associated with an increase of VFI and body fatness at slaughter (Le Dividich et al., 1985), whereas under warm conditions, the VFI is decreased as well as body fatness (Rinaldo and Le Dividich, 1991). However, most of those results were obtained in conventional types of pigs and would not be applicable to lean types of pigs as, for instance, no increase of body fatness under cold temperature was reported recently by Massabie et al. (1996). Furthermore, little information is available to predict VFI and feeding behaviour in pigs exposed to different ambient temperatures (Nienaber et al., 1990, 1996), especially in lean types of pigs. As the chemical composition of the body weight (BW) gain depends directly on the intake of nutrients and their digestive and metabolic utilisation by the animals (Whittemore and Fawcett, 1976), it is of major interest to investigate the way that the ambient temperature affects nutrient utilisation and feeding behaviour in pigs. The aim of the present study was to characterise the effects of exposure to cold or hot temperatures on VFI, feeding behaviour and energy expenditure of group-housed growing pigs over the growing-nishing period. For this latter purpose, the pigs were studied in a respiratory chamber with a subsequent limited groupsize. The present paper will focus on VFI and components of feeding behaviour.

Fig. 1. Cyclic variation of temperature (8C) over each stage of growth in experiments 1 and 2: - - -, adaptation; , measurement.

days at each of the following temperatures: 22, 19, 16, 14 and 128C. Groups 3 and 4 were used to study the effect of hot exposure and then exposed to ambient temperatures varying from 19 to 298C and from 29 to 198C (experiment 2) with three or four consecutive days at each of the following temperatures: 19, 22, 25, 27 and 298C. The 19 and 228C levels were considered as belonging to the thermoneutral zone of group-housed growing pigs. In the second part of the study (experiment 3), two additional groups were studied at an intermediary stage of growth and exposed to the same thermic set as in experiment 1 (group 5) or in experiment 2 (group 6); the initial BW of these pigs was 45 kg on average and the group size was four pigs.

2. Materials and methods

2.2. Animal management 2.1. Experimental design


In the rst part of the study, voluntary feed intake and individual feeding behaviour were measured on four groups of crossbred Pietrain 3 Large White barrows studied over two consecutive stages of growth. The initial BW was around 37 kg at stage 1 and around 63 kg at stage 2. The corresponding group sizes were four and three pigs, respectively. For each group, temperature varied in a cyclic way over 28 days (a thermic set) at each stage of growth (Fig. 1). Groups 1 and 2 were used to study the effect of cold exposure and then exposed to ambient temperatures varying from 22 to 128C and from 12 to 228C (experiment 1) with three or four consecutive At weaning (about 28 days of age), ve to six pigs originating from different litters were selected according to their BW and health status. Three weeks later, at approximately 15 kg BW, they were gathered in a pen. At 25 kg, four of them were moved to the experimental unit. During the whole experiment, the animals were group-housed in a pen (2.3 3 1.6 m) on metal slatted oor upon a small slurry pit. The pen was equipped with a feed dispenser and a drinking station and placed into a respiratory chamber (12 m 3 ). Faeces, urine and water spillages were collected in a slurry pit located below the pen. The pigs were fed ad libitum a cereal based diet offered as pellets, the composition of which is presented in Table 1.

N. Quiniou et al. / Livestock Production Science 63 (2000) 245 253 Table 1 Composition of experimental diet Composition (g / kg) Wheat Barley Corn Soybean meal 454 Wheat bran Oil L-lysine HCl Dicalcium phosphate Calcium carbonate Vitamin and trace mineral mixture Salt Dry matter (DM) Analysed chemical composition (g / kg of DM) Ash Crude protein Crude fat NDF ADF ADL Starch Crude bre Digestible energy (MJ / kg of DM) Metabolizable energy (MJ / kg of DM) Net energy (NE) (MJ / kg of DM)a
a

247

220.0 220.0 200.0 244.5 40.0 10.0 0.5 20.0 10.0 10.0 5.0 876.0

68 208 35 167 53 5 490 42 15.73 15.07 11.24

electronic identication system (antenna) that was activated by earresponders as the animal entered the station. The access to the feed was adapted to the size of animals with a traverse allowing only one animal to enter. The hopper was lled up daily with an amount of feed sufcient to meet the appetite of the group of pigs. The trough was continuously weighed and when the load cell detected it as unsteady, it corresponded to a visit. After each visit, time and amounts of feed at the beginning and at the end of the visit were recorded. Feed consumption per visit was calculated as the difference between the amounts recorded just before and after the visit with an accuracy of 10 g. Ingestion time per visit corresponded to the difference between the time at the end and at the beginning of the visit. If the feeder was detected as unsteady but no difference in feed amount was noted after the visit, it was not recorded. Water was supplied outside the feeder by a nipple drinker to avoid disturbing the feed weight measurements.

2.4. Descriptive components of feeding behaviour


A single meal can occur in different successive visits with short pauses (within-meal intervals) within which the computer can detect a steady weight. In order to allow comparison with the results from other pigs or other studies carried out under various feeding conditions, successive visits were grouped into the same meal by using the so-called meal criterion dened as the maximum length of the within-meal interval between visits. Then, if visits were separated by intervals longer than the meal criterion, they are considered as different meals. The meal criterion was assessed from our data using a log survivorship function as described by Bigelow and Houpt (1988). A sample of 183 data obtained on individual pigs originating from both experiments and exposed to the different temperatures was randomly chosen to calculate the meal criterion. Ninetyve percent of the individual estimated meal criteria were below 2 min; this value was retained for further calculations. For each pig at each level of temperature, at each stage of growth and on each day of measurement, the following behavioural criteria were calculated: the average number of visits per day; average number of

Estimated from digestible energy content (DE, MJ / kg of DM) and chemical components (g / kg of DM) according to the relationship proposed by Noblet et al. (1994): NE 5 0.703 3 DE 1 0.0066 3 crude fat 1 0.002 3 starch 2 0.041 3 crude protein 2 0.0041 3 crude bre (RSD 5 0.18).

Animals had free access to water. The experiment started after one week of adaptation to the respiration chamber environment and the diet. During this period, the temperature was xed at 228C (experiment 1 and group 5 in experiment 3) or 198C (experiment 2 and group 6 in experiment 3). The relative humidity was 70%. The pigs were weighed at the beginning and the end of each thermic set and at the beginning of the 168C level (experiment 1 and group 5 in experiment 3) or the 258C level (experiment 2 and group 6 in experiment 3).

2.3. Equipment
The pen was equipped with a single-space electronic feed dispenser, which consisted of a trough and a hopper connected through a load cell to a computer. The feeding station was provided with an

248

N. Quiniou et al. / Livestock Production Science 63 (2000) 245 253

meals per day; average feed intake per day (g); average total ingestion time per day (min), dened as the total duration of all the visits performed during the day; average total time of feed consumption (min), dened as the sum of ingestion time plus within meal intervals; average rate of feed intake (g / min), dened as the feed intake per day divided by the total ingestion time; average feed intake per meal (g) and the average total ingestion time per meal (min), dened as the total duration of the visits belonging to the same meal. Behavioural criteria were also estimated over the day and the night separately. In fact, over the three or four consecutive days at each temperature level, only the individual feeding behaviour calculated over the two or three last ones were used for further analysis of data. Indeed, during the rst day at each temperature, the ambient temperature decreased or increased by 2 or 38C over 4 h and this day was dedicated to adaptation to the new temperature.

The proportion of diurnal feed intake was calculated for each group at each thermic level as the ratio between the mean diurnal and mean total feed intake. The response of VFI to temperature and BW and their interaction was studied from the combination of all individual data obtained at each temperature level and at each stage of growth in experiment 1 (N 5 120), experiment 2 (N 5 120) and experiment 3 (N 5 56). A covariance analysis was performed with the group within stage of growth as a xed effect and temperature, BW and their interaction as covariates.

3. Results The average daily gain was 980 and 950 g / d in experiments 1 (groups 1 and 2) and 2 (groups 3 and 4), respectively. A problem occurred during fullling the trough at one of the 198C levels studied over the rst stage of growth, both in experiments 1 and 2. Consequently, number of observations reported in Tables 2 and 3 was lower at this temperature. When exposed to 19 and 228C, the VFI of pigs was higher in experiment 2 than in experiment 1 (2.40 vs. 2.31 kg / d on average, respectively); the mean BW was also slightly higher in experiment 2. Then, separated variance analyses were performed on data originating from each experiment. The mean BW over the two stages of growth studied was 47 and 74 kg in experiment 1 and 49 and 76 kg in experiment 2. The increase of BW between stages of growth was associated with a decreased daily number of meals (23.6 and 2 2.1 in experiments 1 and 2, respectively), the sizes of which increased similarly in both experiments (Tables 2 and 3). As a consequence, the VFI increased with increased BW but to a lesser extent in experiment 1 ( 1 0.49 kg / d) than in experiment 2 ( 1 0.63 kg / d). At thermoneutrality, i.e. at 19 and 228C, the data of both experiments were used to assess the relationship between VFI (VFI 19 22 , g / d) and BW (kg). The covariance analysis indicates that the effect of BW on VFI was not linear: VFI 19 22 5 55.36 BW 2 0.251 BW 2 (RSD 5 353) On average, the rate of feed intake was 10 g / min

2.5. Statistical analysis


Data from experiments 1 and 2 were used to characterise the effects of temperature and BW on the components of feeding behaviour. Results of experiment 3 (only one intermediary stage of growth was considered) were introduced in the data set in order to establish an equation for prediction of VFI from temperature and BW: 120, 120 and 56 individual data were obtained in experiments 1, 2 and 3, respectively. Taking into account social facilitation occurring among group-housed pigs (Hsia and Wood-Gush, 1983), components of feeding behaviour were calculated for each group at each temperature level over each stage of growth, as the average of the four (stage 1) or three (stage 2) individual values obtained at this temperature (ascending or descending). When the information was not available for one pig at one of the two thermic levels, then its data were not taken into account at this temperature. From the data obtained in experiments 1 and 2, 34 mean values were calculated in each experiment and used to test the effects of temperature, stage of growth and their interaction through an analysis of variance (GLM, SAS, 1990). The effect of the group was also introduced in the model within each stage of growth.

N. Quiniou et al. / Livestock Production Science 63 (2000) 245 253 Table 2 Effect of cold temperatures and stage of growth on components of feeding behaviour a Temperature (8C) 12 Number of observations Mean body weight (kg) 4 61 14 8 61 per pig 68 10.6 2.46 78 93 33 16 8 61 19 6 61 22 8 61 Stage of growth 1 16 47 2 18 74 10 S*** RSD b

249

Statistical analysis c

Mean components of daily feeding behaviour Number of visits 78 Number of meals 10.7 Feed intake (kg) 2.53 Total ingestion time (min) 75 Total consumption time (min) 100 Rate of feed intake (g / min) 35 Characteristics of the meal Feed intake (g) Consumption time (min)

73 11.5 2.45 69 85 37

72 11.3 2.39 69 84 37

70 11.5 2.22 60 76 38

83 12.9 2.17 73 90 31

61 9.3 2.66 67 85 42

15 1.9 0.3 7 10 5

S***, G*** S*** S*** T **, S*, G** T **, G** S***

272 10.7

262 9.6

236 8.1

228 8

225 7.4

183 7.7

306 9.9

50 1.4

S*** T **, S***, G***

Diurnal feeding behaviour (% of total) Number of meals 65 Feed intake per day 63 Ingestion time 66
a b

66 60 64

65 59 61

66 62 62

68 63 65

66 61 65

66 62 63

5 6 8

G*** G*

Experiment 1, adjusted means. Residual standard deviation. c From analysis of variance with temperature (T ), stage of growth (S), T 3S interaction and group within stage of growth (G) as main effects. Each observation corresponds to the mean of four or three individual values obtained within each group at each stage of growth at each of the two thermic levels (one at 128C) within the experimental scheme designed. Statistical signication: *** P,0.001, ** P,0.01, * P,0.05.

higher at stage 2 than at stage 1 (P,0.001). Taking into account the difference in VFI between stages, it resulted in a comparable total ingestion time at both stages of growth (Tables 2 and 3). The change of ambient temperature was associated with a variation in VFI. In experiment 1 (Table 2), despite a nonsignicant difference among treatments, when the temperature decreased, VFI increased signicantly from 2.20 kg / d at 228C to 2.53 kg / d at 128C. No signicant interaction was found between temperature and BW on this criterion. In experiment 2 (Table 3), the increase of temperature from 19 to 298C induced a signicant decrease in VFI (2.40 vs. 1.82 at 19 and 298C, respectively). Covariance analysis on the data from experiments 1, 2 and 3 indicates that the effects of temperature (T, 8C) and BW (kg) on VFI (g / d) were both quadratic with a signicant interaction between both factors. Between 12 and 298C, the equation was:

VFI 5 2 1264 1 73.6BW 2 0.26BW 2 1 117T 2 2.40T 2 2 0.95T 3 BW (RSD 5 329) The ambient temperature did not inuence the number of meals per day whichever the experiment concerned (Tables 2 and 3). In experiment 1, meal size was not signicantly affected by temperature but the highest (272 g) and the lowest (225 g) values were observed at 12 and 228C, respectively. When temperature increased from 19 to 298C (experiment 2), no signicant effect of temperature was found on meal size (242 g / meal on average), the lowest value being observed at 298C (205 g / meal). In both experiments, the meal duration increased with a decrease in temperature (P,0.01 in experiment 1 and P50.06 in experiment 2). On average, the rate of feed intake was 36 and 39 g / min in experiments 1 and 2, respectively. No effect of temperature was

250

N. Quiniou et al. / Livestock Production Science 63 (2000) 245 253

Table 3 Effect of hot temperatures and stage of growth on components of feeding behaviour a Temperature (8C) 19 Number of observations Mean body weight (kg) Mean components of daily feeding Number of visits Number of meals Feed intake (kg) Total ingestion time (min) Total consumption time (min) Rate of feed intake (g / min) Characteristics of the meal Feed intake (g) Consumption time (min) 6 63 behaviour 63 11.2 2.40 64 81 39 22 8 63 per pig 66 11.3 2.39 63 81 38 25 8 62 27 8 62 29 4 62 Stage of growth 1 16 49 2 18 76 9 S*** RSD b Statistical analysis b

66 9.8 2.30 59 76 40

58 9.9 2.10 55 68 39

50 10.1 1.82 46 56 40

59 11.5 1.89 57 70 34

62 9.4 2.52 58 75 44

7 1.6 0.27 6 7 4

T **, G*** S**, G*** T *, S***, G*** T *** T *** S***, G***

248 8.5

247 8.2

271 8.8

241 7.6

205 6.2

179 6.7

306 9.0

47 1.4

S*** S***, G**

Diurnal feeding behaviour (% of total) Number of meals 69 Feed intake per day 65 Ingestion time 66
a b

68 65 67

61 64 65

55 60 61

55 62 62

60 60 61

63 66 67

5 5 5

T ***, T 3S* S**, G* S**

Experiment 2, adjusted means. See Table 2. Each observation corresponds to the mean of three or four individual values obtained within each group at each stage of growth at each of the two thermic levels (one at 298C) within the experimental scheme designed.

found on the rate of feed intake under hot exposure or under cold exposure. The ingestion time increased with decreased temperatures in experiment 1 (60 and 75 min / d at 22 and 128C, respectively, P,0.05) and it decreased with increased temperatures in experiment 2 (64 and 46 min / d at 19 and 298C, respectively, P,0.001). About 2 / 3 of feed consumption occurred during the day (Tables 2 and 3). This proportion was neither affected by temperature or by stage of growth in experiment 1. Under warm exposure, the diurnal feed intake was proportionally more important at stage 2 (66 vs. 60% at stage 1), but no signicant effect of temperature was observed. As presented in Table 3, the partition of meals between day and night was signicantly inuenced by temperature: 69% of daily meals were performed during the day at 198C but only 55% at 298C. Such a difference was not observed under cold temperatures. In addition, the interaction between temperature and stage of growth was signicant on the diurnal partition of number of meals (Table 3). The decrease of the proportion of diurnal meals between 19 and 298C was more

important during stage 1 (from 68 to 49%) than during stage 2 (from 70 to 62%).

4. Discussion The effect of increased ambient temperature on the VFI in growing-nishing pigs has been widely studied, and in a recent literature review, Le Dividich et al. (1998) reported that the associated decrease of VFI ranged from a minimum value of 40 g / 8C / d to a maximum of 80 g / 8C / d, which depends on many factors including breed, BW range, diet and, to the highest extent, the temperature range. Even if Close (1989) proposed a linear relationship between VFI and temperature, there is some evidence that VFI varies quadraticaly with ambient temperature (Nienaber and Hahn, 1983; the present study), the change in VFI with temperature being as important as the temperature is high. On average between 12 and 298C, the estimated decrease of VFI with each increase of 18C in temperature would be 38, 51 and 49 g / 8C / d on

N. Quiniou et al. / Livestock Production Science 63 (2000) 245 253

251

average in 60 kg pigs according to the equation obtained in the present study or by Nienaber and Hahn (1983) and Close (1989), respectively. Apart from an eventual effect of genotype on appetite, Nienaber and Hahn (1983) and Close (1989) used data obtained either on individually- or pair-housed animals, whereas group-size was four or three pigs in our study. According to Nienaber et al. (1991), four pigs per group would be the minimal number of animals needed to facilitate huddling under cold exposure. Then, the effect of low temperatures on VFI would be markedly increased in smaller groupsize. This hypothesis could explain the differences in slopes between our study and those of Nienaber and Hahn (1983) and Close (1989). However, these slopes are not very far from the decrease in VFI with increased temperature (17288C) observed in more practical conditions, i.e. 5040 g / 8C / d according to Massabie et al. (1996) in 25105 kg group-housed barrows and gilts, respectively. Through the covariance analysis, our study indicates a signicant interaction between BW and temperature on VFI, which was also reported by Close (1989). In practical terms, the interaction found between temperature and BW means that the heavier the pigs are the more they are sensitive to hot ambient temperatures (Fig. 2). Such a result would be consistent with a decrease of the upper critical temperature of pigs with increasing BW as already

Fig. 2. Variation of voluntary feed intake (VFI, g / d) with temperature (T, 8C) and body weight (BW, kg) calculated using the equation: VFI5 21264173.6BW20.26BW 2 1117T22.40T 2 20.95T 3BW.

suggested by Holmes and Close (1977). From the proposed equation, it can be calculated that the decreases in VFI between 19 and 298C were 48 and 77 g / d / 8C in 45 and 75 kg pigs, respectively, while between 22 and 128C, the increases in VFI were 7 and 36 g / d / 8C in 45 and 75 kg pigs, respectively. In fact, these values indicate the highly detrimental effect of high temperatures on heavier pigs and the inability of light pigs to increase their VFI at low temperatures, probably because of their limited gut capacity. According to our results, the daily number of meals remained constant under the temperature range studied, whereas meal size seemed to decrease with increased temperature. These results are in agreement with data obtained in group-housed pigs under hot conditions by Nienaber et al. (1993). However, in individually kept pigs, Nienaber et al. (1990) showed that increased VFI under cold temperature resulted from an increase of daily number of meals, whereas their size and duration were kept constant. Yet, in individually kept pigs, no huddling behaviour can occur to reduce cutaneous heat losses. In such case, a higher meal frequency is associated with higher physical activity that induces a specic heat production. According to our results, under hot exposure, ingestion time per day was reduced as well as the occupation time of the feeding station. But when ingestion time and consumption time obtained in our study are compared, it appears that the non-eating activity in front of the trough averaged 17 min / d when ambient temperature was below 278C, whereas at 27 and 298C, this activity was reduced to 13 and 10 min, respectively. As physical activity is associated with a high heat production in the pigs (Noblet et al., 1993), the lower duration of non-eating activity under hot exposure could be interpreted as an adaptation to high temperatures. The increase of BW inuenced components of feeding behaviour in experiments 1 and 2 as well. As already reported by Auffray and Marcilloux (1980), Bigelow and Houpt (1988), Nienaber et al. (1990) and Labroue et al. (1995) on group-housed or singlehoused pigs, the increase of BW was associated with a decreased number of meals per day, whereas ingestion time remained about constant. Concomitantly, in all studies, the duration of each meal and

252

N. Quiniou et al. / Livestock Production Science 63 (2000) 245 253

the rate of feed intake increased, which resulted in a higher meal size and an increased total feed intake. The rate of feed intake was slightly higher in experiment 2 than in experiment 1, which would be in agreement with Nienaber et al. (1991) who observed a lower rate of feed intake when temperature was 128C below than 48C above the lower critical temperature. Such a difference could be explained by a decrease in the rate of feed intake during the last visits belonging to the same meal when its size increased markedly and almost reached the maximum gut capacity of the pig. The feeding behaviour of conned pigs studied in the present experiment was mainly diurnal as more than 62% of feed intake occurred during the day. This partition is close to the value obtained by Auffray and Marcilloux (1980) and by Bigelow and Houpt (1988) in individually kept pigs (64%). In group-housed pigs, Labroue et al. (1995) reported a higher proportion of VFI during the day (75%). Opposing those studies but in agreement with Nienaber et al. (1990), our results show that the average proportion of diurnal feed intake did not increase markedly with BW. However, the low number of pigs per group may contribute to the constancy of the diurnal partition of VFI. Indeed, it could be supposed that the housing conditions in the present study induced social facilitation as dened by Hsia and Wood-Gush (1983) with or without mild competition, whereas there was competition in the study carried out by Labroue et al. (1995). In addition, the effect of BW on diurnal partition of feed intake seemed to depend on the temperature. Indeed, a higher increase of the proportion of diurnal feed intake was observed between stages 1 and 2 under hot exposure rather than under cold conditions.

moregulation. Under hot temperatures, decreased VFI is inexorably associated with decreased performance. The decrease or increase in feed intake under hot or cold exposure, respectively, occurs mainly through changes in meal size rather than in daily number of meals. Then, the question arises on the mechanisms implied in determination of meal size. It could be limited by short-term thermic effect of feed under hot exposure as internal heat expenditure is reduced in such conditions. Under cold exposure, meal size would rather be determined by stomach capacity than by heat dissipation. However, metabolic control of voluntary feed intake is complex and more knowledge is required to characterise the effect of ambient temperature on the metabolic pathways involved in feeding behaviour.

Acknowledgements The authors gratefully acknowledge A. Roger, F. Le Gouevec and S. Daniel for their technical assistance.

References
Auffray, P., Marcilloux, J.C., 1980. Analyse de la sequence ` alimentaire du porc, du sevrage a letat adulte. Reprod. Nutr. Develop. 20 (5B), 16251632. Bigelow, J.A., Houpt, T.R., 1988. Feeding and drinking patterns in young pigs. Physiology and Behavior 43, 99109. Close, W.H., 1989. The inuence of the thermal environment on the voluntary food intake of pigs. In: Forbes, J.M., Varley, M.A., Lawrence, T.L.J. (Eds.), The Voluntary Food Intake of Pigs. Occasional Publication no. 13-BSAP, pp. 8796. Fuller, M.F., Franklin, M.F., McWilliam, R., Pennie, K., 1995. The response of growing pigs, of different sex and genotype, to dietary energy and protein. Anim. Sci. 60, 291298. Holmes, C.W., Close, W.H., 1977. The inuence of climatic variables on energy metabolism and associated aspects of productivity in the pig. In: Swan, H., Lewis, D. (Eds.), Nutrition and Climatic Environment, pp. 5174. Hsia, L.C., Wood-Gush, D.G.M., 1983. The temporal patterns of food intake and allometric food intake and allomimetic feeding by pigs of different ages. Appl. Anim. Ethol. 11, 271282. Kanis, E., Koops, W.J., 1990. Daily gain, food intake and food efciency in pigs during the growing periods. Anim. Prod. 50, 353364. Labroue, F., Gueblez, R., Meunier-Salaun, M.-C., Sellier, P., 1995. Inuence de la race sur le comportement alimentaire de porcs en croissance eleves en groupe: premiers resultats dune

5. Conclusion The present study clearly indicates that voluntary feed intake depends on body weight and temperature with a marked negative effect of high temperatures in heavier pigs. Under cold temperature, feed intake increased but it seems that light pigs have limited ability to increase VFI when exposed to cold temperatures. Consequently, extra feed intake does not necessarily meet the extra energy demand for ther-

N. Quiniou et al. / Livestock Production Science 63 (2000) 245 253 comparaison Large White-Pietrain. Journees Rech. Porcine Fr. 27, 175182. Le Dividich, J., Desmoulin, B., Dourmad, J.-Y., 1985. Inuence de la temperature ambiante sur les performances du porc en croissance-nition en relation avec le niveau alimentaire. Journees Rech. Porcine Fr. 17, 275282. Le Dividich, J., Noblet, J., Herpin, P., van Milgen, J., Quiniou, N., 1998. Thermoregulation. In: Wiseman, J., Varley, M.A., Chadwick, J.P. (Eds.), Proc. of the 58th Easter School in Agricultural Science, Progress in Pig Science, Nottingham University Press, pp. 229264. Massabie, P., Granier, R., Le Dividich, J., 1996. Inuence de la temperature ambiante sur les performances zootechniques du ` porc a lengrais alimente ad libitum. Journees Rech. Porcine Fr. 28, 189194. Nienaber, J.A., Hahn, G.L. 1983. Performance of growing-nishing swine in response to the thermal environment. ASAE Oral presentation MCR 83-137, St-Joseph, MI, 12 pp. Nienaber, J.A., McDonald, T.P., Hahn, G.L., Chen, Y.R., 1990. Eating dynamics of growing nishing swine. Trans. ASAE 33 (6), 20112018. Nienaber, J.A., McDonald, T.P., Hahn, G.L., Chen, Y.R., 1991.

253

Group feeding behavior of swine. Trans. ASAE 34 (1), 289 294. Nienaber, J.A., Hahn, G.L., Korthals, R.L., McDonald, T.P., 1993. Eating behavior of swine inuenced by environmental temperature. Trans. ASAE 36 (1), 937944. Nienaber, J.A., Hahn, G.L., McDonald, T.P., Korthals, R.L., 1996. Feeding patterns and swine performance in hot environments. Trans. ASAE 39 (1), 195202. Noblet, J., Shi, X.S., Dubois, S., 1993. Energy cost of standing activity in sows. Livest. Prod. Sci. 34, 127136. Noblet, J., Fortune, H., Shi, X.S., Dubois, S., 1994. Prediction of net energy value of feeds for growing pigs. J. Anim. Sci. 72, 344354. Rinaldo, D., Le Dividich, J., 1991. Assessment of optimal temperature for performance and chemical body composition of growing pigs. Livest. Prod. Sci. 29, 6175. S.A.S., 1990. S.A.S. / STAT Users Guide: statistics, Statistical Analysis Systems Institute, Cary, NC (Release 6.07). Whittemore, C.T., Fawcett, R.H., 1976. Theoretical aspects of a exible model to simulate protein and lipid growth in pigs. Anim. Prod. 22, 8796.