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,2

* Adaptation Physiology Group and
and
Animal Nutrition Group, Wageningen Institute of Animal Sciences, Wageningen University, Wageningen, The Netherlands and
and
Institute for Sugar Beet Research, Nutrition Department, Bergen op Zoom, The Netherlands
2 Correspondence:
P.O. Box 338, 6700 AH, Wageningen, The Netherlands (phone: +31 (0) 317 483 120; fax: +31 (0) 317 485 006; E-mail:
Martin.Rijnen{at}12move.nl).
| Abstract |
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Key Words: Circadian Rhythm Feed Intake Metabolizable Energy Physical Activity Pigs Polysaccharides
| Introduction |
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However, reported results on effects of dietary fiber on time spent on physical activity are not very consistent. Some authors report a decrease in physical activity by feeding fiber-rich diets to individually (e.g., Ramonet et al., 1999; 2000b) or group-housed sows (e.g., Brouns et al., 1994; Danielsen and Vestergaard, 2001), whereas others report no effects (e.g., Whittaker et al., 1998; Ramonet et al., 2000a). Moreover, it can be hypothesized that effects of dietary fiber on physical activity are dependent on their botanical origin (Noblet and Le Goff, 2001). Schrama et al. (1998) reported dose response effects of sugar beet pulp silage (SBPS) on physical activity in growing pigs. Dose response studies on the effects of dietary composition on physical activity of sows, however, are lacking.
It can be hypothesized that the level of dietary SBPS affects physical activity of group-housed sows. Furthermore, it can be hypothesized that an effect of dietary SBPS on physical activity of sows is not constant during the day. In the present study, effects of dietary SBPS content and intake of fermentable nonstarch polysaccharides (fNSP) from SBPS on physical activity and heat production in group-housed sows were studied.
| Materials and Methods |
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Twelve groups of six sows were used. The experiment consisted of a 33-d preliminary period followed by a 7-d experimental period. Groups were randomly assigned to one of four experimental diets. These diets differed mainly in starch and NSP content by an exchange of SBPS for tapioca (on DM basis). The four experimental diets contained, on a DM basis, 0, 10, 20, and 30% SBPS, respectively. Sows were group-fed, using a long trough. Sows receiving the 0% SBPS diet were fed at 1.3 times the assumed maintenance energy requirements. Sows fed the other diets received similar amounts of DM. Groups were housed in environmentally controlled respiration chambers (Verstegen et al., 1987b). Sows were exposed to 12 h of light (about 300 lx, from 0700 to 1900) and 12 h of partial darkness (about 10 lx, from 1900 to 0700) to allow video recordings. Details on animals, housing conditions, and diets are reported by Rijnen et al. (2001).
Measurements
For each group, time spent eating was measured using time-lapse video recorders. The recording of eating time started as soon as the sows had access to the feed and stopped when the trough was empty (i.e., observed from video recordings). With these data and measured DMI, feed consumption rate per group was calculated.
Total heat production (HP) was measured at 9-min intervals by determining exchange of oxygen, carbon dioxide, and methane (indirect calorimetry) as described by Verstegen et al. (1987b). These gaseous exchanges were used to calculate HP according to the formula of Brouwer (1965). During the last 6 d of the experimental period, HP was measured continuously.
Behavior of the sows was recorded using time-lapse video recorders during two separate days (i.e., two whole 24-h periods) within the 7-d experimental period. An instantaneous scan sampling technique (as described by Altmann, 1974) was used to analyze the video recordings for behavioral characteristics (Table 1
). The analyses of video recordings for behavioral characteristics were done at 3-min intervals. At these 3-min intervals, the number of sows that exhibited a specific behavior was recorded. With these data, the percentage of time of all sows that was spent on a specific behavior was calculated. The average of each performed behavioral characteristic was calculated for the same 9-min intervals as HP (i.e., average of three sampling moments). The behavioral characteristic "standing" includes standing up, standing, eating, walking, and lying down (Table 1
). Therefore, physical activity is defined as standing up, standing, eating, walking, lying down, plus sitting. Per group and per day (i.e., per each 24-h period), the 9-min data on HP were related to physical activity according to the following equation:
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![]() | [1] |
where HPij = heat production during day period i and 9-min period j, in kJkg-0.75d-1; µ = overall mean; Di = fixed effect of day period i (i = 1 [from 0700 to 2200], 2 [from 2200 to 0700]); Xj = physical activity of sows during 9-min period j, in percentage of time of all sows spent on "standing plus sitting;" ß = regression coefficient of heat production on physical activity; and eij = error term.
Heat production and physical activity exhibit circadian rhythms (e.g., Aschoff et al., 1974; Schrama et al., 1996). The circadian rhythm in HP is only partially accounted for by physical activity, which has been demonstrated in pigs (van der Hel et al., 1984; Henken et al., 1991). Therefore, a fixed effect of day period with two levels was included in Eq. [1]
. The day was divided into a day period from 0700 to 2200, during which time the sows were most active, and a night period from 2200 to 0700, during which the sows were inactive. The increase in HP around feeding time is not fully related to the elevated physical activity (Verstegen et al., 1987a; Noblet et al., 1993). In the calculation according to Eq. [1]
, data around feeding time, from 0800 to 0900, were excluded. These data were omitted in order to avoid possible bias by inclusion of heat increment associated with food ingestion in the relationship between HP and physical activity.
The heat production related to physical activity (AHP) was calculated as follows:
![]() | [2] |
where AHPj = physical activity related heat production during 9-min period j, in kJkg-0.75d-1; Xj = physical activity during 9-min period j of the video recordings, in percentage of time of all sows spent on "standing plus sitting;" b = the estimated regression coefficient from HP on physical activity from Eq. [1]
. Heat production not related to physical activity or resting heat production (RHP) was derived by subtracting AHP from HP. Physical activity-related HP and RHP were calculated for each 9-min period, including the 1-h period around feeding (i.e., from 0800 to 0900).
The energy cost for physical activity (ECact, in kJkg-0.75d-1) was derived from HP on physical activity, as follows:
![]() | [3] |
where b = the estimated regression coefficient from HP on physical activity from Eq. [1]
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Statistical Analysis
Group was the experimental unit. Mean values of HP, AHP, RHP, and the behavioral characteristics were analyzed for the effect of diet by ANOVA. In addition, least squares means were used for treatment comparisons. A contrast method was used to analyze differences between low- and high-fiber diets (i.e., 0% + 10% vs 20% + 30% SBPS). Moreover, HP, AHP, and RHP were analyzed for the effect of diet by linear regression of these traits on the daily intake of fNSP (expressed in gkg-0.75d-1). The SAS software (SAS Inst., Inc., Cary, NC) was used in all statistical evaluations. The data on HP, AHP, and RHP are expressed in kJkg-0.75d-1.
| Results and Discussion |
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The number of groups and animals used in the analyses is given in Table 2
. Six sows were removed at the start of the experimental period because they came into estrus or had leg problems. At the start of the experimental period, average BW was 229 kg (SEM = 12.3). Average parity of sows was 4.1 (SEM = 1.11). During the experimental period, no feed refusals occurred. Consequently, the daily DMI of the groups was similar for the different dietary treatments (P = 0.18; Table 2
) and averaged 38.3 gkg-0.75d-1.
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Studies on the effects of dietary fiber on the behavior of sows have focused mainly on feeding motivation, feeding rate, aggressive behavior, and the occurrence of oral stereotyped activities (as reviewed by Meunier-Salaün et al., 2001). To study these effects, most studies analyze behavior during short periods during the day, mostly related to feeding time (e.g., Braud et al., 1998; Whittaker et al., 1999; Danielsen and Vestergaard, 2001). In the present study, however, only general activity (or postures) was analyzed for during 24-h periods.
In the present study, sows spent on average 163 min/d standing, 14 min/d sitting, 272 min/d lying on breast, and 991 min/d lying on flank (Table 3
). Furthermore, in the present study, dietary composition affected activity level of group-housed sows (P < 0.01; Table 3
). Contrast analysis showed that sows fed 0 or 10% SBPS spent more time standing and "standing + sitting" than sows fed 20 or 30% SBPS (P < 0.01).
In the present study, time spent on physical activity (i.e., "standing plus sitting") averaged 177 min/d, which is low compared with data in literature. In the literature, average time spent on physical activity ranges between 190 min/d (Noblet, 1990) and 363 min/d (Ramonet et al., 1999) for individually housed sows, and between 202 and 435 min/d for group-housed sows (Brouns et al., 1994). In addition, the variation between individual sows in time spent on physical activity is large (e.g., Cariolet and Dantzer, 1984; Noblet et al., 1993). Differences between studies in time spent on activity might be due to housing conditions, feeding level (Terlouw and Lawrence, 1993), physical condition (Cariolet and Dantzer, 1984), and characteristics of the diet (Meunier-Salaün et al., 2001), like dietary fiber content and origin (e.g., Ramonet 1999; 2000a,b).
Reported effects of dietary fiber on physical activity of sows are not very consistent. Due to differences among experiments in breeds, diets, feeding level, recording periods, and housing conditions, it is difficult to look at effects of dietary fiber between studies (Meunier-Salaün et al., 2001). Ramonet et al. (2000a), for example, reported no effect of dietary fiber level (i.e., mixed fiber sources) on standing behavior, whereas Ramonet et al. (1999) reported decreased standing activity with similar diets. Ramonet et al. (2000a), however, used sows housed individually and insulated in respiration chambers, whereas Ramonet et al. (1999) used individual pens. Terlouw and Lawrence (1993) did not find a difference between individually tethered and loose-housed sows in total activity during the day. Furthermore, several authors report a difference in total activity time caused by feeding level; a high feeding level decreased time spent on physical activity (e.g., Halter et al., 1980; Susenbeth and Menke, 1991; Terlouw and Lawrence, 1993). This is important, because in most studies feeding level (kg) is increased because of the decreased dietary energy density when dietary fiber content increases.
Moreover, differences between studies might be related to the level and botanical origin of dietary fiber (Noblet and Le Goff, 2001). Most reported effects of dietary fiber on physical activity during the day have been found using sugar beet pulp as fiber source (e.g., Brouns et al., 1994; Braud et al., 1998; Ramonet et al., 2000b). The decrease in physical activity with higher dietary SBPS content in the present study is similar to the decrease in physical activity with high dietary sugar beet pulp content reported by Brouns et al. (1994) and Ramonet et al. (2000b). In addition, Ramonet et al. (1999) reported a decrease in standing activity of sows that were fed a diet high in mixed fiber sources. And Ramonet et al. (2000b) reported a decrease in standing activity when sows were fed a diet high in wheat bran. Specific components or physicochemical properties of specific dietary fiber sources might be of importance for the effect on behavior and activity (Brouns et al., 1995; Noblet and Le Goff, 2001). Moreover, fermentation characteristics (e.g., fermentation rate) of specific fiber sources in the gastrointestinal tract might be of importance.
Energy Cost of Activity
In sows, the energy cost of activity can be divided into the energy cost of sitting, standing, sitting up, standing up (16.4, 34.1, 35.5, and 49.0 kJkg-0.75 per 100 min, respectively; Kelley et al., 1978), and walking. It is possible that housing conditions or feeding frequency interacts with energy cost of activity, because sitting up and standing up cost more energy than sitting and standing, respectively (Kelley et al., 1978).
In the present study, the energy cost of activity averaged 464 kJkg-0.75d-1 (SEM = 31.0; Table 4
), or 32.2 kJkg-0.75 per 100 min, and did not differ between diets (P = 0.679). The ECact of group-housed sows in the present study is similar to values for individually housed sows reported by Kelley et al. (1978), Noblet et al. (1993), and Ramonet et al. (2000a) (i.e., 34.1, 27.3, and 30 kJkg-0.75 per 100 min, respectively). It can be hypothesized that energy cost of activity in sows does not depend on housing conditions (i.e., individually vs group-housed). Due to our experimental setup, it was not possible to make a distinction between energy cost for different physical activities or movements.
Total Heat Production and Activity-Related Heat Production
Total HP and AHP of the sows are shown in Table 4
. Total HP was calculated for the same 2 d when the videotapes were recorded. Total HP and AHP were affected by diet (P < 0.05), whereas RHP was not (P = 0.331). Among others, Terlouw and Lawrence (1993) reported that a higher feeding level resulted in lower physical activity. In the present study, however, ME intake tended to decrease with increasing dietary SBPS content (P = 0.057; Table 2
), whereas sows fed 0 or 10% SBPS spent more energy on activity than sows fed 20 or 30% SBPS (P = 0.006). The mean AHP of group-housed sows in the present study (56 kJkg-0.75d-1) is similar to the AHP value of 62 kJkg-0.75d-1 reported by Noblet et al. (1993). The small difference between the two values might be due to the higher feeding level in the present study compared to the study of Noblet et al. (1993) since there is an effect of feeding level on time spent on activity (e.g., Halter et al., 1980; Susenbeth and Menke, 1991). Compared to the present study, Ramonet et al. (2000a) reported a higher average value of AHP in individually housed pregnant sows of 105 kJkg-0.75d-1. This might be due to differences in the method of measuring activity between studies (i.e., force sensors and video recordings, respectively).
The mean AHP in the present study represented 12.7% of HP, and ranged from 10.7 to 15.6% among groups. This percentage is in agreement with literature values (e.g., Halter et al., 1980; Verstegen et al., 1987a; Noblet et al., 1993). Ramonet et al. (2000a) reported a higher average percentage of AHP of total HP (i.e., 22%) in individually housed pregnant sows, which might be related to differences in the method of measuring activity between studies. In conclusion, dietary composition altered energy expenditure on physical activity of group-housed sows.
Increased fNSP intake, at similar DMI, decreased the 24-h mean HP (P = 0.039; Table 5
) and tended to decrease the 24-h mean AHP (P = 0.063; Table 5
). Sows tended to spend 2.3 kJ less on physical activity per gram increase of fNSP intake. Schrama et al. (1998) found that growing pigs spent 3.9 kJ less on physical activity per gram of fNSP intake. This difference between animals may be related to body mass (Porzig and Liebenberg, 1977). It can also be due to the high feeding level in growing pigs (Halter et al., 1980; Susenbeth and Menke, 1991; Terlouw and Lawrence, 1993). Moreover, in the present study, video recordings were used for 2 d to measure physical activity, whereas Schrama et al. (1998) used a radar device for 6 d to measure physical activity. Furthermore, in the present study, there were small differences in ME intake between diets (P = 0.057; Table 2
). Feeding level can affect physical activity (e.g., Terlouw and Lawrence, 1993). In the present study, however, the effect of ME intake on AHP could have partially masked the pure effect of fNSP.
Circadian Rhythms
General.
The circadian rhythms of average HP and average AHP of the 0 and 30% SBPS groups are illustrated by Figure 1
. Sows were exposed to 12 h of light (from 0700 to 1900) and 12 h of partial darkness (from 1900 to 0700), and fed once a day (at 0800). Similar to the results reported by Noblet et al. (1993), HP and AHP were highest during eating and decreased until about 12 h after the meal. The first increase in HP and AHP during the afternoon (at 1530) was due to an animal keeper entering the respiration chamber to check health of the sows. The second increase in HP and AHP was due to automatically turning down the lights at 1900. Both peaks were therefore caused by the experimental setup.
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Total Heat Production and Activity-Related Heat Production.
To evaluate the variation in HP and AHP within the day, the day was divided into three parts. The part from 2200 to 0700 was designated as the night period. The night period was part of the partial darkness period. Video recording showed that physical activity was minimal during the night period. This is in agreement with the findings of Cariolet and Dantzer (1984) and Ramonet et al. (2000b), who found that sows had reduced activity during the night period. From 0700 to 2200 (i.e., the remainder of the day) was designated as the day period. The third part was the eating period, from 0800 to 0900, during which time the sows were most active. The linear relationships between fNSP intake and HP, and AHP, are calculated for 24 hthe night period, day period, eating periodand 24 h without the eating period (Table 5
).
Averaged over the whole day (24 h), HP decreased with increasing fNSP intake (P = 0.039). During the night period, fNSP intake did not affect HP (P = 0.509). During the day period, however, HP decreased with increasing fNSP intake (P = 0.006). During eating, HP increased with increasing fNSP intake (P = 0.012). After removing the data of the eating period from the whole day, HP decreased with increasing fNSP intake (P = 0.027).
However, ME intake tended to be lower when SBPS content increased (P = 0.057; Table 2
). Assuming that HP varies by about 0.2 kJ for each kilojoule of ME, HP may be corrected for differences in ME intake (Table 4
). Correcting HP for differences in ME intake changes the relationship between HP and fNSP intake:
![]() | [4] |
It can be concluded that differences in ME intake might accentuate or mask effects of treatments on HP. In the present study, it is unlikely that the differences in ME intake accentuate the effect of fNSP intake on physical activity, because the lower feeding level (i.e., ME intake) with increasing SBPS content would increase physical activity and not diminish it, as found in the present study.
Averaged over the whole day (24 h), AHP tended (P = 0.063) to decrease with increasing fNSP intake. During the night period, fNSP intake did not affect AHP (P = 0.516). During the day period, however, AHP tended (P = 0.062) to decrease with increasing fNSP intake. During eating, AHP tended (P = 0.074) to increase with increasing fNSP intake. After removing the data of the eating period from the whole day, AHP decreased with increasing fNSP intake (P = 0.031). There were no effects of fNSP intake on RHP (data not shown).
About 90% of the decrease in mean HP per gram of fNSP intake (2.62 kJ/g; P = 0.039) was caused by the decrease in mean AHP per gram of fNSP intake (2.31 kJ/g; P = 0.063). During eating, about 60% of the increase in HP per gram of fNSP intake (12.52 kJ/g; P = 0.012) was caused by the increase in AHP per gram of fNSP intake (7.36 kJ/g; P = 0.074).
The observed tendencies for alteration of physical activity by dietary composition is in agreement with other studies with growing pigs (Schrama et al., 1998), gilts (Brouns et al., 1994), and sows (e.g., Robert et al., 1993; Brouns and Edwards, 1994; Ramonet et al., 1999; 2000b). As reviewed by Meunier-Salaün et al. (2001), most effects of fibrous diets on sow behavior were found using sugar beet pulp as fiber source. A decrease in activity level of sows with increasing sugar beet pulp intake might be related to the high water-holding capacity of the diets and delayed gastric emptying and/or specific effects of short chain fatty acids that are produced during fermentation in the hindgut (Meunier-Salaün et al., 2001). Furthermore, the total fermentability and/or fermentation rate related to the botanical origin of dietary fiber, might be of importance for an effect of dietary fiber on behavior of pigs (e.g., feeding motivation).
In conclusion, the effect of fNSP intake on energy expenditure for physical activity was not constant during the day. Sows tended (P = 0.074) to spend more energy on physical activity during eating with increasing fNSP intake, but less during the remainder of the day (i.e., 24 h without eating, P = 0.031).
Physical Activity and Net Energy
The regression coefficients in the relationships between mean AHP and fNSP intake (Table 5
) can be interpreted as the saving effect of physical activity on the NE value of the fNSP fraction. During 24 h, the saving effect of AHP on the NE value of fNSP was 2.3 kJ per g of fNSP intake (P = 0.063; Table 5
). The NE value of the fNSP fraction of a diet or feed ingredient (in kJ/g) can be calculated by correcting HP of pigs for similar digestible nutrient intakes, followed by a regression of fNSP intake on the corrected HP (Rijnen et al., 2001). A second method to calculate the saving effect of physical activity on the NE value of fNSP is the difference in regression coefficients between regression of fNSP intake on HP during the 24-h period and during the night period. This method assumes that sows are not active during the night period. According to this method and the regression coefficients in Table 5
, the saving effect of physical activity on the NE value of fNSP from SBPS was 3.7 kJ/g of fNSP intake. According to the relationship between AHP and fNSP intake (Table 5
) and the calculation as described above, the saving effect of physical activity of sows on the NE value of fNSP from SBPS ranges between 2.3 and 3.7 kJ/g of fNSP intake (i.e., fNSP from SBPS). Assuming a NE value of fNSP from SBPS of 13.4 kJ/g (Rijnen et al., 2001), the energy saving effect of physical activity of sows on the NE value of fNSP from SBPS ranges between 18 and 28% of its NE content.
Feed Consumption Rate
The present study shows that fNSP intake (i.e., fNSP from SBPS) of sows had opposite effects on HP during eating and during the remainder of the day. Therefore, the eating period (from 0800 to 0900) is discussed more closely.
Mean eating time was 16 min (Table 6
), which is in agreement with the mean eating time in individually housed sows (13 min) reported by Noblet et al. (1993). Mean eating time is also similar to reported eating times of sows fed low fiber diets (Braud et al., 1998; Ramonet et al., 1999; 2000b). Despite similar DMI among different treatments in the present study, mean eating time was affected by diet (P = 0.048; Table 6
). Contrast analysis showed that sows fed 0 or 10% SBPS spent less time on eating than sows fed 20 or 30% SBPS (P = 0.009). Several authors reported an increased time spent on eating when dietary fiber content increased (e.g., Brouns et al., 1994; Braud et al., 1998; Ramonet et al., 1999; 2000b).
In the present study, 8.8% of total activity (177 min/d) of the sows was spent on eating. Noblet et al. (1993) reported that sows spent 5.8% of their total activity time on eating. This is in agreement with the sows fed 0 or 10% SBPS in the present study (i.e., 6.0% of total activity time was spent on eating). Sows fed 20 or 30% SBPS spent a larger percentage of their total activity time on eating (i.e., 12.7%), which is in agreement with sows fed high fiber diets (i.e., 14.2%) in the study from Ramonet et al. (1999).
The feed consumption rate in the present study averaged 156 g DM/min or 2.7 g DMkg-0.75min-1. The average feed consumption rate was higher than that reported by Dourmad (1993; 95 g/min), Noblet et al. (1993; 120 g/min), and Brouns et al. (1997; 108 g/min), and similar to that of sows fed low-fiber diets, as reported by Ramonet et al. (1999 and 2000b; 152 g/min and 149 g/min, respectively). In the present study, rate of feed consumption (expressed in g DMkg-0.75min-1) was affected by diet (P = 0.016; Table 6
). On average, feed consumption rate was higher for groups fed 0 or 10% SBPS than for groups fed 20 or 30% SBPS (P = 0.003). This is in agreement with findings of several authors, who reported that feed consumption rate of sows fed a control diet was higher than that of sows fed a diet containing high percentages of sugar beet pulp (Brouns et al., 1997; Braud et al., 1998; Ramonet et al., 2000b). As reviewed by Meunier-Salaün et al. (2001), the low feed consumption rate of diets containing high levels of sugar beet pulp might be due to reduced feeding motivation level, increased mastication time, or lowered palatability due to physical and/or metabolic processes during digestion of sugar beet pulp. Guérin et al. (2001) showed a higher water holding capacity and delayed gastric emptying for diets with sugar beet pulp. Furthermore, differences in feed consumption rates between studies might be due to differences in housing conditions, as suggested by Nielsen (1999). Besides, differences between the present study and other studies might also be due to differences in feed preparation (i.e., mash feed vs pellets or meal).
Finally, it was found in the present study that for each gram increase of fNSP intake (i.e., fNSP from SBPS) the feed consumption rate decreased by 11 g DM/min (P = 0.012) or 0.19 g DMkg-0.75min-1 (P = 0.003). Diet composition affected time spent on eating and feed consumption rate of sows.
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| Footnotes |
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Received for publication July 27, 2002. Accepted for publication September 2, 2002.
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