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ANIMAL PRODUCTION |
Department of Animal Science, University of Minnesota, St. Paul 55108
| Abstract |
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16 sows per treatment combination) was observed on d 1 postweaning, and on d 40 and d 80 of gestation. The percentage of time standing, lying, sitting, feeding, inactive, and performing stereotypic behaviors was determined. Saliva samples were collected to determine cortisol concentrations. Sow BW and backfat depth were determined on d 0, 40, and 80 of gestation, within 24 h of farrowing, and at weaning. An energy and nitrogen digestibility study was conducted using 36 sows assigned to each of the 4 treatment combinations. Over a 24-h period, the sows fed the high-fiber diet spent less time lying (P < 0.05) than the sows fed the control diet. The frequency of feeding did not affect sow behavior measured over a 24-h period. During mealtimes, sows fed the high-fiber diet spent more time feeding (P < 0.05) than sows fed the control diet. Feeding the high-fiber diet did not affect stereotypic behavior measured over 24 h or during mealtimes. Neither diet nor feeding frequency affected salivary cortisol concentration. Sows fed the high-fiber diet gained less BW and lost backfat (P < 0.05) during gestation compared with sows fed the control diet, whereas sows fed once daily gained less BW and lost backfat (P < 0.05) compared with sows fed twice daily. Sows fed the high-fiber diet had fewer pigs born (P < 0.05) compared with sows fed the control diet. Feeding frequency had no effect on size or weight gain of litters. Sows fed the high-fiber diet exhibited lower digestibility of DM, energy, and N (P < 0.05) compared with sows fed the control diet. Feeding a high-fiber diet utilizing soybean hulls or increasing feeding frequency did not enhance the welfare of sows by reducing stereotypic behaviors nor did it improve reproductive performance.
Key Words: feeding frequency fiber gestation sow stereotypic behavior welfare
| INTRODUCTION |
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Feeding high-fiber diets during gestation may improve sow welfare. Including high-fiber ingredients in gestation diets provides an opportunity to offer more feed to the sow, while maintaining daily energy intake at desired levels. Feeding high-fiber diets can influence the welfare and reproductive performance of gestating sows by decreasing stereotypic behavior (Meunier-Salaun et al., 2001
) and increasing the number of piglets born (Grieshop et al., 2001
). Feeding frequency may also positively influence sow reproductive performance (Wittman, 1986
) and behavior of gestating sows (Robert et al., 2002
).
Altering diet composition and feeding management simultaneously may improve welfare and performance of stall-housed sows. Therefore, the objective of this study was to evaluate effects of feeding a high-fiber diet and twice-daily feeding on welfare and performance of gestating sows.
| MATERIALS AND METHODS |
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Experimental Treatments
Dietary treatments consisted of a corn-soybean meal-based diet (control) and a high-fiber diet containing soybean hulls as a fiber source (Table 1
). Frequency treatments consisted of feeding sows once (1x) or twice (2x) daily. Experimental diets and feeding frequencies were provided from 1 d after weaning until d 109 of gestation. Sows were not fed on the day of weaning. Dietary energy and amino acid concentrations were formulated to meet or exceed NRC (1998)
nutrient requirement recommendations for gestating sows with a target BW gain of 40 kg during gestation and an expected litter size of 12 pigs. To provide a daily energy intake of 6,200 kcal of ME/d, sows fed the control diet were offered 1.88 kg of feed/d, whereas sows fed the high-fiber diet were offered 2.19 kg of feed/d. The nutrient composition of the soy hulls used for diet formulation was: 2.09 Mcal of ME/kg (Renteria-Flores, 2003
), 12.1% CP, 0.63% lysine, 0.49% Ca, 0.21% P, 50.0% ADF, and 67.0% NDF (NRC, 1982
). It was assumed that soy hulls provided no vitamins or trace minerals. Body condition scores were evaluated subjectively using a target score of 3, which is approximately 18 to 20 mm of backfat at the last rib (van Heugten, 2000
). Quantity of feed offered to sows in all treatment groups was increased at d 40 and 80 of gestation if sow BCS was below 3. Changes in the amount of feed offered during gestation were recorded. Sows fed 1x received their entire allotment of feed at 0730, whereas sows fed 2x received one-half of their allotment at 0730 and the other half at 1430. Sows fed 2x were housed in a room separate from sows fed 1x such that the second feeding did not influence behavior and activity of sows fed only 1x. From d 109 until farrowing, all sows received 2.25 kg/d of a common lactation diet in meal form (Table 1
). After farrowing and throughout lactation sows were provided ad libitum access to the same diet.
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16/treatment combination) were observed using time-lapse video recording for 24 h on the day after weaning, d 40, and d 80 of gestation. Behavior was analyzed using The Observer software (Version 3.0; Noldus Information Technology, Leesburg, VA). Videos were scored 20 min/h for 24 h using 4 intervals of 5 min throughout each hour (e.g., 1200 to 1205, 1215 to 1220, 1230 to 1235, and 1245 to 1250). Postures evaluated included duration of time spent lying, sitting, or standing. Activities recorded included feeding, sham chewing, bar biting, and nosing the feeder or floor. Sham chewing, bar biting, and nosing the floor and feeder were combined to form one category of stereotypic behaviors. Due to the placement of the video recorder, it was not possible to recognize feed in the feeder, but it was clear when feed dropped into the feeder. Therefore, after the feed was dropped, a sow was considered to be feeding from the time her head was observed in the feeder until she moved at least one whole step back from the feeder. When the sow was not involved in any activity, she was recorded as nonactive.
Stress Physiology Measurements
Saliva was collected from focal sows on the day after weaning, and on d 42 and 82 of gestation using a salivette with a cotton swab (Sarstedt, Aktiengesellschaft and Co., Numbrecht, Germany), which was attached to a length of stiff wire. Sows were allowed to chew the device until the swab became saturated with saliva. Samples were collected at approximately 1300 each collection day. On the day after weaning, saliva samples were taken during video recording. However, all other samples were collected after video recording was complete. Saliva was harvested by centrifugation and was frozen at 20°C within 2 h of collection. Saliva samples were analyzed for cortisol concentration using a RIA technique (Coat-a-Count TKCO, Diagnostic Products Corporation, Los Angeles, CA) as described by Anil et al. (2005)
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Energy and Nitrogen Digestibility Determinations
On d 100 of gestation, 36 sows with an average BW of 219.0 kg (range: 191.6 to 256.1 kg) were used in a nutrient digestibility trial to evaluate effects of diet and frequency of feeding on energy and nitrogen digestibility. Only sows in parity 3 or greater were used for the trial. The average parity for these sows was 3.6 (range: 3 to 5). Sows remained in their respective housing environments during the digestibility trial. The digestibility trial was conducted over 6 wk beginning in July, using 3 contemporary groups of sows. A contemporary group was defined as sows due to farrow within 1 wk of each other. Sows were fed their respective experimental diets containing 0.25% chromic oxide. The experimental diets were fed for a 7-d adjustment period followed by a 3-d collection period. Fecal samples were collected twice daily at 0800 and 1600, pooled, placed in plastic bags, and frozen at 20°C. At the end of collection, feces were dried in a force-draft oven (60°C) for 2 d and then weighed. Dried feces were ground through a 1-mm screen and frozen until subsequent nitrogen and gross energy determinations could be performed. Total tract DM digestibility was determined by chromic oxide analysis using the procedure of Fenton and Fenton (1979)
. Nitrogen content of diets and feces was determined by the combustion method using a Leco TruSpec analyzer (Leco Corp., St. Joseph, MI). Gross energy of feed and feces was determined by bomb calorimetry (Parr 1281 Bomb Calorimeter, Parr Instrument Co., Moline, IL). Blood was collected from 6 sows per dietary treatment via jugular venipuncture before the morning feeding on the third day of the collection period for serum urea nitrogen analysis. Following collection, serum was harvested after centrifugation and frozen. Serum urea nitrogen was analyzed using photospectrometry (Biotron Diagnostics, Hemet, CA).
Performance Measurements
Sow performance measurements were recorded during gestation and the subsequent lactation. Sows were weighed and backfat measurements were recorded on d 0, 40, and 80 of gestation, within 24 h of farrowing, and at weaning. Backfat was measured at the last rib using ultrasound (Renco Corp., Minneapolis, MN). After farrowing, litter size at birth and weaning were recorded. Litter size was equalized within diet and frequency treatments to achieve a target of 10 pigs per litter. Sows were fed twice daily during lactation to ensure ad libitum access to feed. Feed not consumed was weighed on d 7 and 14 of lactation, and at weaning, and was subtracted from the total amount of feed offered to determine feed disappearance for each period. Feed wastage was observed subjectively throughout the experiment and was determined not to be a problem. After weaning, sows were moved into gestation crates for breeding and the number of sows that returned to estrus within 8 d was recorded.
Statistical Analyses
Data collected repeatedly throughout a sows gestation and lactation were analyzed by least squares AN-OVA with repeated measures in time using the GLM procedure of SAS (SAS Inst. Inc., Cary, NC). The initial statistical model included effects of diet, feeding frequency, parity group, farrowing group, and all possible interactions. Parity group 1 described sows in parities 1 or 2, whereas parity group 2 described sows in parity 3 and greater. Whenever the 2-, 3-, and 4-factor interactions were not significant (P > 0.05), these factors were pooled into the residual error term. However, interactions with time and the 2-factor interaction of diet and feeding frequency remained in the model regardless of level of significance. Energy and nitrogen digestibility were analyzed by least squares ANOVA. The statistical model included effects of diet, feeding frequency, contemporary group, and all appropriate 2-way interactions. Chi-square was used to analyze the number of sows that returned to estrus within 8 d. All means are reported as least squares means.
| RESULTS AND DISCUSSION |
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Sow Behavior
Over a 24-h observation period, sows fed the high-fiber diet behaved differently (P < 0.05) compared with sows fed the control diet at different stages of gestation (Table 2
). At weaning, behavior was unaffected by dietary or feeding frequency treatments. On d 40 and 80 of gestation, sows fed the high-fiber diet spent a greater percentage of time sitting and a smaller percentage of time lying over a 24-h observation period compared with control-fed sows (P < 0.05). The sows fed the high-fiber diet also spent a greater percentage of time feeding on d 40 and 80 of gestation in agreement with Robert et al. (2002)
who fed high-fiber diets to gilts. During feeding, sows must stand. Those animals fed the high-fiber diet spent more time eating than sows fed the control diet, which may explain some of the increased time spent standing, but this should not be considered undesirable. Stereotypic behaviors and nonactive time were not affected by dietary treatment at any stage of gestation. Feeding sows once or twice daily had no effect on postures or noneating behaviors measured over a 24-h observation period throughout gestation. However, at d 40 and 80 of gestation, sows fed 2x spent a greater percentage of time feeding (P < 0.05) than sows fed 1x. This was not expected because sows fed 2x received a similar amount of feed as sows fed 1x, but in 2 meals instead of 1. Presumably, sows spent some time licking and nosing the feeder trough at the end of a meal before stepping away from the feeder. This activity would have been recorded as feeding 2 times each day for sows being fed 2x compared with only once for sows fed 1x.
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Several factors may have converged to prevent the high-fiber diet from influencing stereotypic behavior of sows. Stereotypic behaviors likely result from frustrated feeding motivation (Lawrence and Terlouw, 1993
). Feeding motivation in gestating sows seems to be driven by a physiological need for energy and a need for gut fill (Whittaker et al., 1998
). Expression of stereotypic behaviors is minimized when both needs are satisfied (Bergeron and Gonyou, 1997
; Whittaker et al., 1998
). Possibly, the physiological needs of our sows for energy was satisfied as evidenced by their lactation performance but the small increase in quantity of high fiber diet offered was insufficient to satisfy the sows needs for gut fill.
Alternatively, the effects of high-fiber diets on sow behavior may have been modulated by body condition. In contrast to their previous work with thinner females (Ramonet et al., 1999
), Ramonet et al. (2000)
reported no differences in the amount of time sows spent standing when fed a high-fiber diet compared with a control diet. Cariolet and Dantzer (1984)
observed that sows in ideal body condition (BCS = 3) spent more time standing than did overconditioned sows, and that thin sows tended to be hyperactive. In the current study, the target subjective BCS was 3. Fifty-eight percent of sows assigned to the high-fiber diet received increases in daily feed allowance during the study, which meant these sows were thin. This may have increased the stereotypic behaviors for these sows and masked any possible positive effects of dietary fiber on reduction of these behaviors.
Finally, age of the sows in our study may have masked beneficial effects of dietary fiber. Incidence of stereotypic behaviors increases with advancing parity of sows (Lawrence and Terlouw, 1993
; van der Peet-Schwering et al., 2003
) and the efficacy of high-fiber diets in redirecting stereotypic behaviors is reduced (van der Peet-Schwering et al., 2003
) due to channeling of frustrated feeding behavior toward stereotypic activities (Lawrence and Terlouw, 1993
). High-fiber diets can reduce stereotypic behaviors in mature sows especially when fed at levels to support positive BW balance (Ramonet et al., 1999
). All sows in our experiment were in at least their second parity with many in their third and fourth parities. Sows fed the high-fiber diet gained less BW during gestation than control sows. Possibly, the age of our sows and the relatively small gestation BW gain of sows fed the high-fiber diet combined to block positive effects of the high-fiber diet on expression of stereotypic behavior.
At d 40 and 80 of gestation, diet had no effect on postures of sows measured around feeding time (Table 3
). During the 3-h feeding period, sows fed the high-fiber diet spent a greater percentage of time eating compared with control sows (P < 0.05). This is likely due to the greater amount of feed offered to sows on the high-fiber diet and the fibrous nature of the diet. Stereotypic and noneating behaviors were unaffected by diet.
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At d 40 and 80 of gestation, sows fed 1x performed stereotypic behaviors a greater percentage of observation time (P < 0.05) and spent a lower percentage of the observation time inactive around mealtimes compared with sows fed 2x. On the surface, it appears that these results conflict with those reported by Robert et al. (2002)
who found that gilts fed one meal per day spent a smaller percentage of time performing stereotypic behaviors around mealtime than gilts fed twice per day. However, actual time spent performing stereotypic behaviors around mealtime for sows fed 1x (122 min on d 40 and 91 min on d 80) was less than sows fed 2x (200 min on d 40 and 149 min on d 80). Sows fed 1x were inactive during the afternoon period when sows fed 2x were eating. We theorized that more frequent feeding would provide the sow more feeding bouts each day that would direct a larger portion of the sows daily time budget to productive activities such as feeding. Twice-daily feeding did increase the amount of time sows engaged in feeding behaviors; however, it also encouraged sows to spend more time engaged in stereotypic behaviors. Ingestion of feed stimulates the performance of stereotypic behaviors in restricted-fed sows (Terlouw et al., 1993
); such behaviors peak soon after feeding (Robert et al., 1993
).
Sow behavior patterns changed as gestation progressed regardless of treatment. On d 40 of gestation, all sows showed increases in percentage of time standing and decreases in percentage of time lying compared with d 0 and 80 of gestation. Sows also spent a greater percentage of time performing stereotypic behaviors on d 40 of gestation compared with d 0 and 80 of gestation. Similarly, Anil et al. (2005)
reported increased standing, decreased lying, and increased stereotypic behaviors measured at d 56 of gestation compared with d 5 postweaning and d 108 of gestation. Sekiguchi and Koketsu (2004)
observed greater incidence of vacuum chewing for sows in the second trimester of gestation compared with the first trimester and greater bar biting than sows in late pregnancy. Time spent standing decreased after d 80 of gestation. Zonderland et al. (2004)
suggested that adaptation to housing and management conditions occurs in the first 6 wk of gestation, which limits expression of stereotypic behaviors. They also theorized that the decline in stereotypic and substrate-directed behaviors after d 84 of gestation is related to a corresponding decline in time spent standing.
Stress Physiology
Neither diet nor feeding frequency had any effect on salivary cortisol concentration of gestating sows (Table 2
). Similarly, McGlone and Fullwood (2001)
noticed no effect of feeding high-fiber diets on plasma cortisol concentration of pregnant gilts. We did observe an important effect of time (P < 0.05) on cortisol concentration regardless of treatment. Cortisol concentration was highest on d 0 and decreased throughout gestation. Anil et al. (2005)
reported similar results for stall-housed sows. If both salivary cortisol (Greenwood and Shutt, 1992
) and stereotypic behaviors (Broom, 1983
) are indicators of stress in livestock, it is not clear why these 2 response criteria followed different patterns throughout gestation.
Nutrient Digestibility
We observed effects of contemporary group (P < 0.05) on daily feed intake during the digestibility study. Likewise, an interaction between feeding frequency treatments and contemporary group (P < 0.05) was observed for feed intake because a greater proportion of sows in the first group fed 1x had their feed intake increased to correct poor body condition compared with sows fed 1 x in the second and third groups. These group effects appear to be due to random chance. We observed no interactions between dietary treatments and contemporary group for DM, gross energy, or nitrogen digestibility.
As expected, sows fed the high-fiber diet had greater ADFI and DMI (P < 0.05), because these females were offered more feed in an attempt to meet their energy requirements compared with sows fed the control diet (Table 4
). Feeding the high-fiber diet also reduced DM digestibility (P < 0.05) in agreement with Pond et al. (1985)
. Feeding frequency affected (P < 0.05) ADFI, DM intake, and GE intake. This unintended effect resulted because 10 out of 17 sows assigned to the 1x treatment had their daily feed amounts increased in midgestation due to poor body condition compared with only 8 of 19 sows assigned to the 2x treatment.
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Sow Performance
Sows fed the high-fiber diet consumed more feed and DE (P < 0.05) than control sows from d 40 through 109 of gestation (Table 5
). An interaction between diet and feeding frequency treatments (P < 0.05) was observed for ADFI (2.25, 2.08, 2.54, and 2.60 kg for control-1x, control-2x, high-fiber-1x, and high-fiber-2x, respectively) and daily DE intake (6,958, 6,318, 7,131, and 7,202 kcal for control-1x, control-2x, high-fiber-1x, and high-fiber-2x, respectively) from d 80 through 109 of gestation.
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Sow BW was similar at breeding, but on d 40 of gestation, sows fed 2x weighed more (P < 0.05) than sows fed 1x. This difference was maintained throughout the remainder of gestation and throughout lactation. At breeding and d 40 of gestation, backfat depth was similar between 1x and 2x treatment groups. By d 80 of gestation, backfat depth was lower for sows fed once daily (P < 0.05); this difference was maintained throughout the remainder of gestation and the subsequent lactation period. No time by dietary or feeding frequency treatment interactions were observed for any measure of sow performance.
We did not expect less BW gain or backfat loss for the sows fed the high-fiber diet because feed allowance was calculated to provide sows with similar levels of energy and other nutrients. However, after d 40 of gestation, 38% of sows fed the control diet and 58% of sows fed the high-fiber diet were offered increased feed allowances. Evidently, the actual energy density of experimental diets was less than calculated creating the need to increase daily feed intake. The variance of actual from predicted energy density was greater for the high-fiber diet as evidenced by the lower gestation BW gain in sows fed the high-fiber diet compared with the control diet. The sows fed the high-fiber diet consumed more feed and digestible energy than control sows but this increase was not sufficient to support pregnancy BW and backfat gains observed in control sows.
Lower BW gain and backfat loss of sows fed the high-fiber diet may be due to behavior of the animals. Sows fed the high-fiber diet spent more time standing and less time lying compared with sows fed the control diet. Noblet et al. (1993)
reported that sows expend increased amounts of energy due to standing (3.56 kcal/min), and it is important to consider the activity level of sows when determining energy requirements. At d 40 and 80, sows fed the high-fiber diet in this experiment spent more time in upright postures (standing plus sitting) and had increased activity (feeding plus stereotypic behaviors) compared with control sows. At d 40, sows fed high fiber would have used 1,666 kcal/d for standing and sitting compared with control sows, which used 1,379 kcal/d. Similarly at d 80 of gestation, high-fiber sows expended 1,413 kcal/d for standing and sitting compared with 1,122 kcal/d for control sows. This increased energy need for standing and sitting contributed to the lower BW gain of sows fed the high-fiber diet.
The greater gestational BW gain of 2x sows compared with 1x sows is puzzling. Sows fed 2x consumed similar amounts of feed but slightly less DE and still gained 8.4 kg more BW than sows fed 1x. One might conclude that sows fed 2x used dietary energy more efficiently than sows fed 1x. However, energy digestibility of diets was not influenced by feeding frequency, similar to the results reported by Sharma et al. (1973)
with young pigs. Although sows fed 2x were housed in a separate room from sows fed 1x, the room temperatures and housing conditions were similar. A plausible explanation for this disparity in sow BW gain is not evident to us. Further investigations are needed to determine if this is a real or spurious result.
Wean-to-estrus interval of sows was unaffected by treatment. Similarly, percentage of sows that expressed estrus within 8 d after weaning was not affected by treatments (89.6 for control-1x, 92.1 for control-2x, 82.7 for high fiber-1x, and 90.2 for high fiber-2x;
2 = 2.95; df = 3).
Litter Performance
Sows fed the control diet had increased total pigs born, pigs born alive, and after cross-fostering (P < 0.05) compared with sows fed the high-fiber diet (Table 6
). There was no effect of diet on litter weight or piglet BW. Feeding frequency had no effect on any litter performance traits, similar to the results of Wittman (1986)
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In addition to timing, source of fiber may be important in eliciting an increase in litter size. Fibrous feed ingredients such as corn cobs (Matte et al., 1994
), oat hulls (Mroz et al., 1986
), alfalfa haylage (Hagen, 1988
), and wheat straw (Ewan et al., 1996
) have supported increases in litter size at farrowing. In contrast, Nelson et al. (1992)
reported a reduction in litter size when sows were fed soybean hulls from breeding to farrowing. Likewise, Renteria-Flores (2003)
found no improvement in litter size when soybean hulls were offered to sows beginning 2 d postmating to d 109 of gestation.
The high-fiber diet and feeding management practice imposed in this experiment were not effective in improving the welfare and reproductive performance of pregnant sows housed in stalls. Some researchers have reported beneficial effects of dietary fiber on sow welfare and reproductive performance whereas others have reported no beneficial effects. The variable response of sows to dietary fiber suggests that numerous factors influence the efficacy of dietary fiber. A more thorough understanding of these factors will enhance the utility of high-fiber diets for commercial pork production.
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| Footnotes |
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2 This research was funded in part by the Minnesota Agricultural Experiment Station. ![]()
3 Present address: North Carolina State University, Dept. of Animal Science, Box 7621, Raleigh, NC 27695. ![]()
4 Corresponding author: johnstlj{at}morris.umn.edu
Received for publication May 25, 2005. Accepted for publication November 21, 2005.
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