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Effects of exposure to high ambient temperature and dietary protein level on performance of multiparous lactating sows

1

D. Renaudeau*, N. Quiniou†, and J. Noblet* ,2

*Station de Recherches Porcines—Institut National de la Recherche Agronomique, 35590 Saint Gilles, France and †Institut Technique du Porc, 35650 Le Rheu, France

ABSTRACT: Fifty-nine multiparous Large White × Landrace sows were used to determine the effects of high ambient temperature and level of dietary heat increment on lactation and reproductive performance. During a 28-d lactation and the 14-d postweaning pe- riod, ambient temperature was maintained constant at 20 or 29°C. Experimental diets fed during lactation were a control diet (NP; 17.6% crude protein) and two low-protein diets obtained by reduction of CP level (LP; 14.2% CP) and both reduction of CP and addition of 4% fat (LPF; 15.2% CP); the NE:ME ratio was 74.3, 75.6, and 75.8% for NP, LP, and LPF diets, respectively. All diets provided 0.82 g of digestible lysine/MJ of NE, and ratios between essential amino acids and lysine were above recommendations. During the ad libitum period (i.e., between d 7 and 27 of lactation), ADFI and NE intake decreased, respectively, from 7.63 to 4.22 kg and 82.0 to 43.3 MJ of NE when the temperature increased from 20 to 29°C. Exposure to 29°C reduced litter BW gain (2,152 vs 2,914 g/d) and increased lactation BW

loss (34 vs 16 kg) but increased postweaning BW gain

(2 vs 4 kg). Expressed per kilogram of feed intake, water intake increased from 4.0 to 7.5 L between 20 and 29°C. Compared with the NP diet, low-CP diets (LP or LPF) maintained performance and reduced total

N excretion (22.5%) in lactating sows at thermoneu-

trality and attenuated the negative effects of high tem- perature on NE intake and BW loss (40.9 vs 47.9 MJ

of NE/d and 41 vs 30 kg for NP and LP + LPF diets,

respectively). Temperature and diet composition did not affect the reproductive performances, and the mean weaning-to-estrus interval was 4.9 d. The respiratory rate and skin, udder, and rectal temperatures increased markedly at 29°C (105 vs 33 breaths per min and 37.8 vs 36.1°C, 38.9 vs 38.2°C, and 39.5 vs 38.8°C, respectively) without any effect of diet. In conclusion, low heat incre- ment diets (i.e., with reduced CP/NE ratio) did not affect lactation performance of sows at thermoneutrality and attenuated the effects of high ambient temperature on energy intake and BW loss. At any ambient tempera- ture, N excretion can be markedly reduced.

Key Words: Dietary Protein, Feed Intake, Heat Stress, Lactation, Sows

2001 American Society of Animal Science. All rights reserved.

J. Anim. Sci. 2001. 79:1240–1249

Introduction

During the last decade, selection for improved pro- lificacy of sows has resulted in increased litter size at weaning and a concomitant increase of the sows’ nutrient requirements. Simultaneously, selection for leaner pigs at slaughter weight was associated with a decrease of body reserves and voluntary feed intake. Consequently,

1 The authors gratefully acknowledge Degussa-Hu¨ ls (Hanau, Ger- many) and Ajinomoto Eurolysine (Paris, France) for their financial support; Ajinomoto Eurolysine (Paris, France) for measurement of amino acid contents in feeds; Institut Technique du Porc (Paris, France) for the grant of D. R.; J. Van Milgen for critical evaluation of the manuscript; and P. Bodinier, S. Daniel, S. Dubois, J. Gauthier, and H. Renoult for their technical assistance. 2 Correspondence: E-mail: noblet@st-gilles.rennes.inra.fr. Received June 19, 2000. Accepted January 5, 2001.

the nutritional deficit of lactating sows has been accentu- ated with subsequent excessive weight loss during lacta- tion and extended weaning-to-estrus interval or success- ful mating (Whittemore, 1996). Apart from genetic selec- tion, high ambient temperature is one of the many factors that may influence the voluntary feed intake of lactating sows (O’Grady et al., 1985). In summer, lactat- ing sows are frequently exposed to high ambient temper- atures and their feed intake is reduced (Black et al., 1993; Prunier et al., 1997; Quiniou and Noblet, 1999). This decreases heat production due to the thermic effect

of feed (TEF). The reduction of energy intake is more

important than the change in energy requirement due

to decreased milk yield, which results in an accentuated

nutritional deficit. Performance of sows exposed to high ambient temper- atures can be maintained by increasing heat loss to the environment (McGlone et al., 1988). An alternative ap- proach is to reduce heat production of the sow. The TEF

1240

High temperature and diet composition

1241

of digestible fat, starch, protein, and dietary fiber aver-

ages 10, 18, 42, and 42% of the ME content, respectively (Noblet et al., 1994). Therefore, diets with reduced crude protein (CP) level and(or) fat addition result in lower heat production (Noblet et al., 1987; Le Bellego et al., 1999) and seem to be better tolerated under hot climatic conditions (Stahly et al., 1979). The objective of our study was to evaluate the effects of diets with reduced TEF on performance and feeding behavior of lactating sows and performance of piglets when exposed to high or con- ventional ambient temperatures. The present paper will focus on performance of sows.

Materials and Methods

Experimental Design. Fifty-nine multiparous, cross-

bred Large White × Landrace sows divided into 11 groups

of four to six animals were used in the experiment. Two

farrowing rooms were used for the experiment, one set

at 20°C (thermoneutral) and one set at 29°C (hot). Each

group of sows was maintained either at 20°C or at 29°C during the 28-d lactation period and the subsequent 14-

d postweaning period. The ambient temperature was

maintained constant over the 24-h period. Within each group, sows were allocated to three dietary treatments (one or two sows per treatment) in order to obtain, for the total experiment, similar parity number and body condition and BW at farrowing for the six treatments. The three experimental diets differed in their NE:ME ratios; differences were achieved by a reduction of the CP level (LP for low protein level) or both reduction of CP and addition of fat (LPF for low-protein and fat- added diet); these diets were compared to a normal CP

level diet (NP). The three diets supplied the same levels

of

lysine (0.82 g digestible lysine/MJ of NE), and levels

of

digestible essential amino acids relative to digestible

lysine were similar and higher than values recom- mended by Dourmad et al. (1991). Mineral and vitamin contents met or exceeded the recommendations of INRA (1989). Composition, chemical characteristics, and nutri-

tional values of the diets are given in Table 1. Care and use of animals met the requirement of the certificate

of Authorization to Experiment on Living Animals, no.

04739 (certified by the French Ministry of Agriculture

to J. Noblet).

Animal Management. In order to standardize body con- dition at farrowing, pregnancy feed allowance was calcu- lated for each sow according to the model proposed by Dourmad et al. (1997) and described previously by Quin- iou and Noblet (1999). The gestation diet contained 13.0 MJ of DE/kg, 13% CP, and 0.55% lysine. Fourteen days before farrowing, the animals were

moved to the farrowing rooms and kept at 24°C with a photoperiod of 14 h of artificial light (0830 to 2230) and

a minimum ventilation rate of 25 m 3 /h per sow. Within

each temperature, sows were allotted to the three dietary treatments according to BW and backfat thickness on d 100 of gestation. Farrowing room conditions and the feeding plan at the beginning of lactation have been

Table 1. Composition of experimental diets

Item

NP

LP

LPF

Ingredient, g/kg Wheat Corn Soybean meal Wheat bran Sugar beet molasses Soy oil L-Lysine HCl DL-Methionine L-Threonine L-Tryptophan L-Isoleucine L-Valine Dicalcium phosphate Calcium carbonate Salt Vitamins and trace minerals mixture

350

403

364

353

406

366

207

91

129

30

30

30

20

20

20

 

40

1.7

5.5

5.3

0.4

1.5

1.7

0.9

2.5

2.7

0.6

0.6

1.3

1.4

2.2

2.3

16

16

16

11

11

11

5

5

5

5

5

5

Chemical composition, % (as fed)

DM

87.3

87.4

88.1

Ash

5.1

5.0

5.5

Crude protein

17.6

14.2

15.2

Crude fat

2.5

2.6

6.1

Starch

43.2

49.5

44.7

Crude fiber NDF ADF ADL Lysine Methionine + cystine Threonine Tryptophan Isoleucine Leucine Valine Histidine Phenylalanine

2.2

2.0

1.9

9.3

9.4

8.9

2.6

2.3

2.4

0.4

0.4

0.4

0.96

0.95

1.04

0.60

0.60

0.64

0.69

0.67

0.74

0.20

0.20

0.21

0.70

0.61

0.66

1.40

1.07

1.15

0.82

0.80

0.84

0.45

0.33

0.35

0.82

0.60

0.65

Digestibility coefficients, % b

DM

89.9

89.5

89.6

N

90.0

88.9

90.6

Energy

91.7

90.9

90.9

Energy value, MJ/kg bc Digestible energy Metabolizable energy Net energy

14.7

14.4

15.2

14.2

14.0

14.8

10.5

10.6

11.2

NE/ME, % Digestible lysine, g/MJ of NE d

74.3

75.6

75.8

0.82

0.82

0.82

a NP = normal protein, LP = low protein, and LPF = low protein and added fat. b Values measured on nonpregnant adult sows (four measurements per diet). c Adjusted for measured DM on the pooled samples. ME was esti- mated from DE content and the ME/DE ratio was assumed to be equal to those measured in growing pigs fed the same diets (96.2, 97.2, and 97.2% for NP, LP, and LPF diets, respectively; Lebellego et al., unpublished data). Net energy was estimated from measured digestible energy content (MJ/kg DM) and chemical component (g/ kg DM) according to Noblet et al. (1994). d Estimated from analyzed lysine content and estimated digestibil- ity coefficient of lysine calculated by additivity from characteristics of raw materials according to Eurolysine and ITCF (1995) tables.

described by Quiniou and Noblet (1999). Litter size was standardized to 12 pigs by cross-fostering within 48 h after birth. When the number of available piglets was too low, piglets from nonexperimental sows were fostered.

1242

Experimental temperature was established on d 1 (i.e., 24 h after the mean farrowing day) over a 4-h period. Water was supplied for ad libitum consumption to the sows from a low-pressure nipple drinker connected to a 55-L graduated water tank. Creep feed (17.3 MJ of GE/

kg, 21.4% CP) was offered to the pigs after d 21 following farrowing. The day before weaning (d 27), refusals were collected at 1600 in order to weigh the sows with an empty digestive tract on the following morning. After weaning, sows stayed in the farrowing room at the same temperature for 14 d to record postweaning reproductive performance. During this period, they received 3.0 kg/d

of the gestation diet and were checked visually twice a

day for estrus using an aerosol reproducing the odor of the boar (Boarmate, ANTEC, France). Measurements. Live weight and backfat thickness were measured at farrowing, at weaning (d 28), and

d 14 after weaning. Backfat thickness was determined

ultrasonically 65 mm from the midline at the last rib (P2 site). Piglets were individually weighed at birth,

every 7 d during lactation, at weaning, and at d 14 after weaning. Every morning, refusals were removed be- tween 0830 and 0900 and new feed was immediately offered as pellets. Average daily feed intake and water consumption were determined as the difference between feed allowance and refusals collected on the next morn- ing. The weaning-to-estrus interval was determined vi- sually (i.e., immobilization while pressing on the back) and confirmed by blood progesterone tests on blood sam- ples taken on d 8 and d 13 after weaning. Nutrient digestibility and energy value measurements

of experimental diets were carried out in a Latin square

design during 10-d periods of excreta collection using three nongravid sows fed at maintenance as described by Noblet and Shi (1993). Body temperatures were measured every Monday morning from the Monday before farrowing to the Mon- day of the 2nd wk after weaning. The ambient tempera-

ture was established on Friday (i.e., the day after the mean farrowing day), so temperatures were measured

at 4, 3, 10, 17, 24, 31 and 38 d of exposure. The respira-

tory rate was measured on the same days, but only dur- ing lactation and only for some sows (n = 18). A digital thermometer and a type K probe were used to measure

the temperatures on the back (i.e., at the P2 position), on the mammary gland, and on an intermediate point between the back and the udder (i.e., flank). A digital thermometer was used to measure rectal temperatures. To determine respiratory rate, flank movements were counted during one minute between 0900 and 1100, but only for resting sows (Quiniou and Noblet, 1999). Chemical Analyses. Feed and feces were analyzed for moisture, ash, and fat according to AOAC (1990) meth- ods, for N according to the Dumas method, and for crude fiber and cell wall components according to Van Soest and Wine (1967). Gross energy was measured using an adiabatic bomb calorimeter. Total amino acid contents

of the diets were analyzed at the Ajinomoto Eurolysine

laboratory (Amiens, France) using ion-exchange chroma-

Renaudeau et al.

tography, except for tryptophan, which was analyzed using high performance liquid chromatography. Nitro- gen in urine was measured on fresh material and urinary energy content was obtained after freeze-drying 50 mL in small polyethylene bags. Concentration of blood pro- gesterone was measured by radioimmunoassays; sows were considered to be in estrus when blood concentration of progesterone exceeded 3 ng/mL (Martinat-Botte et al., 1996).

Calculations and Statistical Analyses. Digestibility coef-

ficients (%) and DE contents (MJ/kg) of experimental diets were calculated according to routine procedures (Noblet and Shi, 1993). Nongravid sows used for digest- ibility measurements had close to zero N retention; ME values obtained under these conditions underestimate the ME value of similar diets fed to lactating sows, which retained about half of the dietary N. Therefore, the ME content (MJ/kg) was estimated from DE content mea- sured in the sows and the ME:DE ratio measured on the same diets in growing pigs (96.2, 97.2, and 97.2% for NP, LP, and LPF diets; Lebellego et al., unpublished data). The NE content (MJ/kg) was estimated from DE content and chemical composition of diet (g/kg MS) ac- cording to the equation proposed by Noblet et al. (1994). Chemical composition of lactation empty BW loss (g/d) was estimated from variations in BW and P2 backfat thickness using the equations proposed by Dourmad et al. (1997). 3 Lactation N loss from body reserves was assessed from estimated mobilization of body protein, assuming a N content of 160 g/kg in body protein. Nitro- gen from creep feed (Ncf) intake was calculated from creep feed intake per litter over the last week of lactation. Nitrogen output in milk (Nm) was estimated for all lacta- tion periods from total N retained by the litter (Re- naudeau and Noblet, unpublished data); Ncf and Nm were assumed to be used by suckling piglets with effi- ciencies of 90 and 87%, respectively (Noblet and Etienne, 1986). Fecal N was estimated using the digestibility coef- ficient of N for each experimental diet. Urinary N was calculated as the difference between N supplied by feed and body reserves and N output in milk and feces. The effects of ambient temperature, diet composition, and their interaction on results were tested through analysis of variance (General Linear Models procedure, SAS Inst. Inc., Cary, NC). The effect of group of sows was tested within the effect of ambient temperature. Contrasts were used to compare the NP diet with the low-CP diets (LP and LPF). The effect of stage of lacta- tion on feed intake was analyzed as repeated measures

3 Lipids (kg) = −26.4 (4.5) + 0.221 (0.030) × empty BW + 1.331 (0.140) × P2 (RSD = 6.1); Protein (kg) = 2.28 (2.22) + 0.178 (0.017) × empty BW 0.333 (0.067) × P2 (RSD = 1.9); Water (kg) = 23.6 (4.4)

+ 0.551 (0.031) × empty BW 0.919 (0.135) × P2 (RSD = 5.2); Ash

(kg) = 0.58 (0.61) + 0.037 (0.005) × empty BW 0.081 (0.018) × P2 (RSD = 0.5); Energy (MJ) = −1,075 (159) + 13.67 (1.12) × empty BW

+ 45.98 (4.93) × P2 (RSD = 208). Empty BW (kg) = a × BW 1.013 (kg),

with a = 0.912 at farrowing and a = 0.905 at weaning. P2 = P2 backfat

thickness (mm).

High temperature and diet composition

Table 2. Effects of ambient temperature and diet composition on litter growth and daily consumption of sows during lactation (adjusted means)

1243

 

20°C

29°C

Statistical

Item

NP a

LP a

LPF a

NP

LP

LPF

RSD b

analysis c

No. of sows Parity Duration of lactation, d Litter size Litter weight gain, g/d

10

9

9

11

11

9

2.6

2.8

3.0

2.9

3.0

3.2

0.8

G*

28.3

28.4

29.0

28.1

27.9

27.8

0.7

T**, G**

10.8

10.6

10.8

10.8

10.8

10.1

1.0

G*

2,933

2,872

2,938

2,150

2,242

2,064

494

T***, G*

Intake from farrowing to weaning Feed, g Water, L

Intake from d 7 to 27 Feed, g DE, MJ ME, MJ NE, MJ Water, L Water/feed, L/kg

6,707

6,507

6,727

3,560

4,054

3,862

790

T***

25.5

25.4

30.9

28.7

25.1

31.3

7.6

G*, D†

7,736

7,541

7,630

3,874

4,513

4,264

910

T***

112.5

108.5

116.7

56.3

65.0

65.2

13.4

T***

108.2

105.4

113.4

54.2

63.1

63.3

12.9

T***

80.4

79.7

85.9

40.2

47.7

48.0

9.5

T***, D†

29.0

29.2

34.2

31.5

27.1

34.3

8.9

3.7

3.8

4.5

8.2

6.1

8.2

1.8

T***

a NP = normal protein diet, LP = low-protein diet, and LPF = low-protein and added fat diet. b Residual standard deviation. c From analysis of variance including the effects of ambient temperature (T), diet composition (D), interaction between temperature and diet composition (T × D), the linear contrast between NP and LP + LPF diets (C), and the effect of group of sows within temperature (G). Statistical significance: ***P < 0.001, **P < 0.01, *P < 0.05, †P < 0.10.

with temperature as main effect (Repeated procedure, SAS Inst. Inc.). Data for body temperatures and respiratory rate were analyzed with a multifactorial analysis of variance (split plot) taking into account the effects of temperature, diet, duration of exposure to ambient temperature, and inter- actions. Data from d 4 of exposure were not taken into account for testing the interaction between ambient tem- perature and duration of exposure because experimental temperature was set on d 1 after farrowing. The effects of location of measurement (only for body temperatures) were also taken into account in the preceding model without the effect of duration of exposure. Number of sows coming into estrus before and 5 d after weaning were compared using a chi-square test (SAS Inst. Inc.).

Results

The sow parity number was 2.9 on average, and the mean litter size during lactation was 10.6 (Table 2). The lactation length was lower for one group of sows kept at 29°C with a subsequent shorter lactation at 29°C than at 20°C (27.9 vs 28.6 d); no difference was observed between dietary treatments. Compared with an exposure to 20°C, exposure to 29°C depressed ADFI during the whole lactation period and over the ad libitum intake period (i.e., between d 7 and 27) (3,825 vs 6,647 and 4,217 vs 7,635 g/d, respectively; Table 2). The effect was similar whether based on daily ME or on NE intakes. At 29°C, even though dietary treatments had no effect (P > 0.10) on ADFI and daily NE intakes, those values were similar (P = 0.36) for the LP and LPF diets and numerically higher than for NP diet (47.9 vs 40.2 MJ of NE/d). However, at 20°C, daily

ME or NE intakes were similar (P > 0.10) for NP and LP diets and lower than for the LPF diet (80.0 vs 85.9 MJ of NE/d). As a consequence, NE intake was less depressed by exposure to high ambient temperature when sows were fed the LP diet (minus 40%) than when they were fed the NP diet (minus 50%); the LPF diet was intermediate (minus 44%). Ambient temperature had no influence (P > 0.10) on daily water intake either over the total lactation or during the ad libitum intake period (Table 2). This resulted in a water:feed intake ratio that was higher at 29°C (7.5 vs 4.0 L/kg at 20°C, P < 0.001). At the beginning of lactation period, the sows were restrictively fed, so that ADFI increased similarly at both ambient temperatures until d 5 (Figure 1). At 29°C, between d 5 and 6, ADFI decreased by 677 g/d (P < 0.001). After d 6, ADFI increased progressively with the advancement of lactation (+752 and +587 g/d between

the advancement of lactation ( + 752 and + 587 g/d between Figure 1 . Daily

Figure 1. Daily feed intake during the lactation period (20°C = ; 29°C = ×).

1244

d 6 and 26 at 20 and 29°C, respectively; P < 0.05) (Fig- ure 1). Lactation BW loss was lower at 20°C than at 29°C (16 vs 34 kg, P < 0.001, Table 3), but it was not affected by dietary treatment. However, despite a tendency of interaction between temperature and diet (P = 0.10), sows kept at 29°C and fed the low-CP diets lost less BW than those fed the NP diet (30 vs 41 kg). Consequently, when temperature increased from 20 to 29°C, the BW loss variation was more pronounced with the NP diet (+ 160%) than with the LP or LPF diets (+80%). At far- rowing, backfat thickness tended (P < 0.10) to be lower at 29°C than at 20°C (19.8 vs 21.4 mm) and variation over lactation was not affected by temperature and di- etary treatment (3.7 mm on average). Over the post- weaning period (i.e., between weaning and d 14 after weaning), sows’ BW increased by 2 kg at 29°C and de- creased by 4 kg at 20°C; backfat thickness change was not affected by ambient temperature (+0.5 mm on aver- age). The latter criteria were not affected by dietary treatment during the previous lactation. The empty BW loss was higher (P < 0.001) at 29°C than at 20°C (1,260 and 633 g/d, respectively), in connec- tion with higher (P < 0.001) water, lipid, protein, ash, and energy mobilization (Table 4). The water, protein, and ash losses at 29°C tended to be higher (P < 0.10) for the NP diet than for the average of the LP and LPF diets (680, 214, and 43 g/d vs 503, 157, and 31 g/d, respec- tively). Effects of ambient temperature and diet composition on the sows’ N balance are presented in Table 5. The increase of temperature from 20 to 29°C resulted in a decrease (P < 0.001) of N intake (44%) and total excreted

Renaudeau et al.

N (35%) and an increase (P < 0.001) of N mobilized

from body reserves (+130%). Reduction of dietary CP level markedly decreased (P < 0.001) the amount of N excreted by the sow for both temperatures (minus 7% for each 1% reduction of dietary CP). When the sows and

their litters were considered together and contribution of N from body reserves was taken into account, N excreted, expressed as a percentage of total N supplied (i.e., N from feed + N from creep feed + N from body reserves), was lower (P < 0.05) for the LP diet (62%) than for the NP diet (65%); the corresponding value for the LPF diet was intermediate (63%). Reproductive performances were measured for a total

of 58 sows because one sow on the NP treatment was

slaughtered after weaning. The weaning-to-estrus inter- val was unaffected by temperature or diet composition (Figure 2). Most sows (49 out of 58) came into estrus within 5 d following weaning; one sow did not show estrus signs before d 14 after weaning. The mean wean- ing-to-estrus interval was 4.9 d for the 57 sows dis- playing estrus before d 14 after weaning. Ovulation was

confirmed in the 57 sows by a blood sample concentration

of progesterone higher than 3 ng/mL.

For technical reasons, BT were not recorded for one group of sows at d 4 and 24 of exposure and for two groups of sows at d 31 and 38 of exposure; 18 sows were measured for respiratory rate at all stages of exposure. Over the lactation period, respiratory rate and back, flank, udder, and rectal temperatures increased (P < 0.01) at 29°C without effect (P > 0.10) of dietary treat- ment (Table 6). Because no effect of location was found for values obtained on the back and the flank, skin tem- perature calculated as the mean of back and flank tem-

Table 3. Effects of ambient temperature and diet composition on body weight and backfat thickness changes of sows over the 28-d lactation and postweaning period (adjusted means)

 

20°C

29°C

Statistical

Item

NP a

LP a

LPF a

NP

LP

LPF

RSD b

analysis c

No. of sows

10

9

9

11

11

9

Body weight, kg At farrowing At weaning At d 14 after weaning Lactation BW change Postweaning BW change d

Backfat thickness (P2), mm At farrowing At weaning At d 14 after weaning Lactation P2 change Postweaning P2 change d

260

258

264

268

264

257

15

G***

244

242

247

227

235

227

18

T**, G***

238

239

243

233

236

231

16

G***

16

15

18

41

29

31

12

T***

6

3

4

1

1

5

5

T***, G*

21.3

22.0

21.0

19.1

20.5

19.7

3.5

T†, G*

17.6

18.5

18.5

14.6

16.4

16.1

3.4

T**, G*

17.9

19.5

19.0

14.7

16.6

16.9

3.4

T**, G†

3.7

3.5

2.5

4.5

4.0

3.6

2.2

0.3

1.0

0.5

0.1

0.3

0.8

1.1

a NP = normal protein diet, LP = low-protein diet, and LPF = low-protein and added fat diet. b Residual standard deviation. c From analysis of variance including the effects of ambient temperature (T), diet composition (D), interac- tion between temperature and diet composition (T × D), the linear contrast between NP and LP + LPF diets (C), and the effect of group of sows within temperature (G). Statistical significance: ***P < 0.001, **P < 0.01, *P < 0.05, †P < 0.10. d Two sows were culled between weaning and d 14 after weaning (in treatment NP at 29°C).

High temperature and diet composition

Table 4. Effects of ambient temperature and diet composition on composition of sows’ BW loss over a 28-d lactation (adjusted means) a

1245

 

20°C

29°C

Statistical

Item

NP b

LP b

LPF b

NP

LP

LPF

RSD c

analysis d

No. of sows Empty body weight loss, g/d Backfat thickness loss, (P2) mm/d

Mobilization of chemical components Water, g/d Lipid, g/d Protein, g/d Ash, g/d Energy, MJ/d

10

9

9

11

11

9

627

596

676

1,503

1,107

1,179

419

T***

0.13

0.12

0.09

0.16

0.14

0.13

0.08

223

216

292

680

477

529

196

T***, C†

316

295

266

547

438

435

178

T**

67

65

91

214

149

166

62

T***, C†

12

12

18

43

29

33

13

T***, C†

14.7

13.8

13.3

27.9

21.8

22.1

8.5

T***

a Empty BW and chemical composition were calculated at farrowing and weaning using equations of Dourmad et al. (1997; see text). b NP = normal protein diet, LP = low-protein diet, and LPF = low-protein and added fat diet. c Residual standard deviation. d From analysis of variance including the effects of ambient temperature (T), diet composition (D), interaction between temperature and diet composition (T × D), the linear contrast between NP and LP + LPF diets (C), and the effect of group of sows within temperature (G). Statistical significance: ***P < 0.001, **P < 0.01, *P < 0.05, †P < 0.10.

peratures was used as a single criterion to compare with udder and rectal temperatures. At 20°C, skin tempera- ture was lower (P < 0.001) than udder temperature (36.2 vs 38.2°C), and both temperatures were lower than rectal temperature (38.8°C). At 29°C, all these values were higher (P < 0.001) than at 20°C, but the differences be- tween locations were reduced: the gradient between skin and rectal temperatures was lower (1.6 vs 2.6°C at 20°C, P < 0.001), whereas the gradient between udder and rectal temperatures remained constant (0.6°C). During the lactation period, no effect (P = 0.15) of duration of

exposure was found on skin temperature. The respira- tory rate increased from 33 to 105 breaths/min when the ambient temperature changed from 20 to 29°C, but it was not affected by dietary treatment. Both skin and rectal temperatures decreased after weaning (i.e., between d 24 and 38 of exposure) (Figure 3). The skin temperature reduction was greater at 20°C than at 29°C (2.9 vs 1°C between d 24 and 38, P < 0.01), whereas rectal temperature change was the same for both temperatures (0.9°C on average). Consequently, after weaning the gradient between skin and rectal tem-

Table 5. Effects of ambient temperature and diet composition on nitrogen balance over a 28-d lactation (adjusted means)

 

20°C

29°C

Statistical

 

Item

NP a

LP a

LPF a

NP

LP

LPF

RSD b

analysis c

No. of sows

10

9

9

11

11

9

N

supplied, g

From feed (Ni)

6,137 i

4,811 j

5,377 j

3,269 k

2,950 k

2,985 k

662

T***, D**, C*, G† T***, C* T**, G**

From

body reserves (Nr) d

305 i

299 i

422 i

968 k

665 l

738 l

285

From

creep feed (Ncf)

61 i

65 i

41 i

115 j

92 j

78 j

47

N

output, g

In milk (Nm) e Excreted (N urine + N feces) Sows f

2,369 i

2,300 i

2,431 i

1,725 j

1,671 j

1,530 j

424

T***, G†

4,073 i

2,809 j

3,369 k

2,512 l

1,945 m

2,193 m

527

T***,

D***, C†

 

Piglets g Total Total, % (Ni + Ncf + Nr) h

297 i

289 i

303 i

223 i

214 i

196 i

52

T***, G†

4,370 i

3,098 j

3,672 k

2,736 l

2,159 m

2,389 m

513

T***,

D***, C†

67 i

60 j

63 ij

63 ij

58 j

63 ij

7

D*

a NP = normal protein diet, LP = low-protein diet, and LPF = low-protein and added fat diet. b Residual standard deviation. c From analysis of variance including the effects of ambient temperature (T), diet composition (D), interaction between temperature and diet composition (T × D), the linear contrast between NP and LP + LPF diets (C), and the effect of group of sows within temperature (G). Statistical significance: ***P < 0.001, **P < 0.01, *P < 0.05, †P < 0.10. d Nr = body protein mobilized during lactation/6.25; body protein calculated from equation of Dourmad et al. (1997); see Table 4. e Nm = (total N retained by piglets Ncf × 0.90) × efficiency of N milk utilization (i.e., 1.14; Noblet and Etienne, 1986). Litter’s N retention was calculated according to the comparative slaughter technique (Renaudeau and Noblet, unpublished data). f N feces = (1 digestibility coefficient of nitrogen) × N in feed, N urine = Ni + Nr Nm N feces. g N excreted = Nm × (1 efficiency of N milk utilization). h N total excreted expressed as a percentage of total N supply (Ni + Ncf + Nr). i,j,k,l,m Effect of temperature and diet composition. Within a row, means not followed by the same superscript differ (P < 0.05).

1246

Renaudeau et al.

1246 Renaudeau et al. Figure 2 . Effect of ambient temperature and diet on number of

Figure 2. Effect of ambient temperature and diet on number of sows displaying estrus after weaning (white portion = estrus 5 d after weaning and black portion = estrus > 5 d after weaning). Mean weaning to estrus inter- val (5.2 d at 20°C and 4.9 d at 29°C) was calculated without one sow not displaying estrus before d 14 after weaning (in LP treatment at 29°C). NP, normal protein; LP, low protein; LPF, low protein and added fat.

peratures was 2.1°C at 20°C but remained the same as during lactation at 29°C (1.7°C).

Discussion

Heat Dissipation at High Ambient Temperatures in Lactating Sows

Independently of the ambient temperature during lac- tation, skin and rectal temperatures increased after far-

tation, skin and rectal temperatures increased after far- Figure 3 . Effect of ambient temperature (20

Figure 3. Effect of ambient temperature (20°C = ; 29°C = ×) and duration of exposure on rectal (———) and skin (– – – –) temperatures).

rowing, remained constant thereafter during lactation, and decreased after weaning. Similar variations in rectal temperature over the successive periods of the reproduc- tive cycle were described by Prunier et al. (1997). This observation can be compared with the direct relationship between heat production and energy supply in lactating sows (Noblet and Etienne, 1987). We can then suppose that elevated body temperatures during lactation com- pared with those in the prefarrowing and postweaning periods is related to the increase of metabolic heat pro- duction due to increased feed intake and milk synthesis.

Table 6. Effects of ambient temperature and diet composition on body temperature and respiratory rate in sows (adjusted means)

Duration of exposure, d

Temperature, °C a

Item

4

3

10

17

24

31

38

20

29

RSD b

Statistical analysis c

Observations, n

54

59

59

59

54

49

50

111

120

Body temperature, °C

Back

35.5

d

36.8

e

37.0

e

36.8

e

36.9

e

35.6

d

35.1

f

36.1

i

37.6

i

0.8

T**, E**, T × E**, A** T**, E**, T × E**, A** T**, E**, T × E**, A** T**, E**, T × E**, A**

Flank

35.8

d

36.9

e

37.2

e

37.0

e

37.2

e

35.7

d

35.1

f

36.2

i

37.9

i

0.7

Udder

36.7

d

38.4

e

38.6

e

38.5

e

38.6

e

37.1

d

35.5

f

38.2

h

38.9

h

0.6

Rectum

38.1

d

39.2

e

39.2

e

39.1

e

39.1

e

38.3

d

38.2

d

38.8

g

39.5

g

0.5

Gradient of body temperature, °C

 

Rectum back Rectum flank Rectum udder

2.6

de

2.4

e

2.3

e

2.2

e

2.2

e

2.7

d

3.0

f

2.7

1.8

0.7

T**, E**, T × E**, A** T**, E**, T × E**, A** T**, E**, T × E**, A**

2.3

d

2.3

d

2.1

d

2.1

d

1.9

d

2.6

e

3.1

f

2.6

1.6

0.7

1.4

d

0.7

e

0.6

e

0.6

e

0.5

e

1.2

f

2.7

f

0.6

0.6

0.4

Respiratory rate, breaths per min

 

Observations, n

18

18

18

18

36

36

Rate

66

70

66

74

33

105

17

T**, A*

a Means calculated only with lactation data. b Residual standard deviation. c From multifactorial analysis of variance (split plot) including the effects of ambient temperature (T), diet composition (D), duration of exposure to the temperature (E), interactions, and effect of animal (A). Statistical significance: **P < 0.01, *P < 0.05. d,e,f Within each line, adjusted mean values with different superscripts are different (P < 0.05). Data from d 4 of exposure were not taken into account for testing the interaction between temperature and duration of exposure because experimental temperature was set on d 1 after farrowing. g,h,i Effect of site of measurement (back, flank, udder, or rectum) was taken account in the model (see footnote b). Within each column, adjusted mean values with different superscripts are significant (P < 0.05, RSD = 1.1).

High temperature and diet composition

1247

One of the effects of high ambient temperatures is an increase of skin, udder, and rectal temperatures; our results (+0.18, +0.08, and +0.08°C per extra degree of temperature, respectively) are in agreement with those of Quiniou and Noblet (1999) (+0.15, +0.05, and +0.11°C between 22 and 29°C). The elevation of peripheral tem- perature can be explained by an increase of blood flow in the skin’s vessels for dissipation of body heat. Moreover, elevated ambient temperature induced a rise in respira- tory rate (+8.0 breaths/min/°C) similar to values reported in other studies (+7.5, +7.1, and +8.9 breaths/min/°C, according to Schoenherr et al., 1989, Lorschy et al., 1991, and Quiniou and Noblet, 1999, respectively). Similarly, in cattle given ad libitum access to feed, Hahn (1999) reported an increase of respiratory rate (+4.3 breaths min 1 °C 1 ) between 18 and 32°C. Cattle have more sweat glands than pigs, and the contribution of the lungs to evaporative heat losses is less important. In contrast, taking into account the low cutaneous evapo- rative capacities in pigs, an important rise in respiratory rate is required in pigs for an effective increase of evapo- rative heat losses. All these processes correspond to a short-term adaptation to hot climatic conditions via in- creased heat dissipation. The reduction in the gradient between rectal and skin temperatures (i.e., 1°C) ob- served in the present experiment when ambient temper- ature increased from 20 to 29°C is in agreement with results of Quiniou and Noblet (1999). No significant effect of duration of exposure at 29°C was observed on skin and rectal temperatures and respi- ratory rate in our study. This means that, in agreement with Quiniou and Noblet (1999), lactating sows adapt quite quickly to hot temperatures (i.e., within 3 d, Figure 3), with no more adaptation on a long-term basis. At a higher temperature (i.e., 32°C) and with lower productiv- ity sows, Schoenherr et al. (1989) reported a reduced rectal temperature and respiratory rate in lactating sows as the lactation progressed. This suggests a tendency for long-term adaptation in heat-stressed sows with an increase of efficiency of evaporative heat loss. The dis- crepancy between both sets of data could be explained by differences in sows’ productivity and its change over lactation.

Thermogenesis at High Ambient Temperatures in Lactating Sows

The effects of high ambient temperature on voluntary feed and energy intakes have been well described in the literature. As reviewed by Quiniou and Noblet (1999), in multiparous and mixed-parity sows, each single de- gree of rise in farrowing house temperature causes a decrease in ADFI ranging between 100 and more than 200 g/d (or 1.3 to more than 2.6 MJ of ME/d); this vari- ability can be explained by differences in lactation length, parity number, milk production, and tempera- ture range considered. Indeed, in contrast to what has been considered in the review of Black et al. (1993), the effect of temperature on ADFI in lactating sows becomes

more pronounced at high ambient temperature (Stans- bury et al., 1987; Quiniou and Noblet, 1999). Similar conclusions have been obtained in growing pigs (Giles and Black, 1991; Quiniou et al., 2000a). Our value (331 g d 1 °C 1 and 4 MJ of ME d 1 °C 1 between 20 and 29°C) is higher than the average value suggested by Black et al. (1993) (170 g d 1 °C 1 and 2.4 MJ of ME d 1 °C 1 ) but is quite similar to those obtained in the recent study of Johnston et al (1999) (289 g d 1 °C 1 and 3.9 MJ of ME d 1 °C 1 between 19 and 27°C) or calculated from the equa- tion proposed by Quiniou and Noblet (1999). It can also be suggested that the higher negative effect of hot tem- peratures on ADFI of lactating sows observed in recent studies can be related to the higher production level at thermoneutrality and the associated higher rate of heat production in highly productive sows with subsequent higher susceptibility to heat stress. At 29°C, sows were restrictively fed at the beginning of the lactation period and seemed not to be affected by high ambient temperature; consequently, ADFI linearly increased until d 5. However, ADFI was significantly reduced from d 5 to 6 (Figure 1), suggesting that sows compensated for previous feed restriction on d 5 and they had a delayed response to ambient temperature. Similar results have not been reported in the literature.

Body Composition in Heat-Stressed Lactating Sows

The BW loss measured at thermoneutrality in our study was considerably higher than the value reported

by Johnston et al. (1999) in multiparous sows (560 vs

60 g/d). In that study, ADFI was rather similar to ours

(i.e., 6,375 vs 6,647 g/d) but litter weight gain and litter size were lower (2,110 vs 2,914 g/d and 9.9 vs 10.7 piglets, respectively). This suggests that the ad libitum supply of high-quality diets under favorable climatic conditions was unable to meet requirements of highly productive sows. The BW loss contained 46% lipids, 38% water, and 12% protein; lipids then represented the major source of energy from body reserves (86%). For multiparous sows studied over a 21-d lactation, Quiniou and Noblet (1999) reported similar values of body nutrient change (15% and 33%, respectively, for protein and lipid content in BW loss) in sows fed diets comparable to ours (i.e., 0.70 g/MJ of ME). Moreover, Sauber et al. (1998) showed that more protein was mobilized when the lysine:ME ratio was below 0.85 g/MJ of ME. However, reduction of energy supply at constant daily protein supply was associated only with accentuated body fat mobilization (Noblet and Etienne, 1987). Consequently, BW loss com- position depends to a large extent on the ratio between protein (i.e., amino acids) and energy in the feed. Between 20 and 29°C, BW loss increased greatly (34 vs

16 kg), whereas backfat thickness loss increased slightly.

The lack of significant effect of temperature on backfat thickness loss may be due to a great variability between sows (CV = 60%) and partly by a lower backfat thickness before farrowing in sows exposed to 29°C.

1248

Although postweaning ADFI was similar for all sows (i.e., 3 kg/d), those kept at 20°C lost BW, whereas the BW of those at 29°C increased during the 14-d postweaning period. Such a difference was also reported by Quiniou et al. (2000b) for sows exposed either at 20 or 26°C during lactation and 21°C after weaning. Hughes (1993) re- ported that feed restriction (3 kg/d) during lactation in- duced a BW gain (+260 g/d), whereas sows fed to appetite (6 kg/d) lost BW (1,111 g/d) over the 7-d period following weaning. This suggests that the postweaning BW change is dependent on the difference in feeding levels between the lactation and the postweaning period. Under conven- tional conditions (i.e., high level during lactation and lower after weaning), BW loss is in fact associated with an important reduction in gut fill and weight of viscera (e.g., gut and liver), whereas lean and adipose tissues start to be reconstituted. Conversely, in sows fed restric- tively during lactation, BW change during the postwean- ing period corresponds to a reduction of mammary gland, positive gain in body reserve tissues, and minor changes in weight of viscera (Ziopoulos et al., 1983; King and Dove, 1987).

Renaudeau et al.

with a subsequent marked reduction of N excretion. Our results and those of Everts and Dekker (1994) and Dour- mad et al. (1998) indicate that N excretion can be reduced by 7% for each 1% reduction of dietary CP. In agreement with Tilton et al. (1995), there was a tendency for higher NE intake for the fat-supplemented diets (i.e., 8%) with- out reduction of ADFI and BW loss.

Low Heat Increment Diets at High Ambient Temperatures

Results of the present study suggest that the negative effects of high ambient temperature on energy intake and mobilization of body reserves are attenuated when

low-CP diets are fed. There are few studies on the effect

of

low-CP diets on performance of lactating sows exposed

to

high ambient temperatures. In response to a reduced

protein supply (i.e., 17 vs 14%), Quiniou and Noblet (1999) reported no improvement in performance of lac-

tating sows kept at 29°C. In that study, the lack of inter- action between temperature and diet was attributed to the low number of observations and(or) the lower sup- plies of sulfur amino acids and tryptophan expressed as

a percentage of lysine in the 14% CP diet. However,

Lynch (1989) observed an increase of feed consumption (+700 g/d) in multiparous, lactating sows fed a low-CP diet (14 vs 20%) at 28°C. Moreover, increasing the energy

concentration of the sows’ diet by addition of fat resulted in an increase of daily ME intake without any effect on BW loss in hot conditions (Schoenherr et al., 1989; Quiniou et al., 2000b). However, according to our results, the combination of reduced CP level and increased fat content (i.e., with 4% of vegetable oil) attenuated the negative effects of high temperature on overall perfor- mance of lactating sows, including BW loss. These re- sults would confirm the benefit of reducing metabolic heat production under high ambient temperature with

a low heat increment diet (low CP with or without fat).

Implications

The present experiment demonstrates that the reduc- tion of CP level in a wheat-corn-soybean meal diet com- bined with supplementation of crystalline lysine and other essential amino acids (methionine, threonine, iso- leucine, tryptophan, and valine) allows the main- tainance of performance and reduction in N excretion in lactating sows kept under thermoneutral conditions. The detrimental effect of exposure to high ambient tempera- ture on feed intake and lactation performance can be attenuated by using diets with low thermic effect of feed.

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