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Impact of diet deprivation and subsequent overallowance during gestation on lactation performance of primiparous sows

Impact of diet deprivation and subsequent overallowance during gestation on lactation performance... Impact of diet deprivation and subsequent overallowance during gestation on lactation performance of primiparous sows ,2 † ‡ § # Chantal Farmer,* Kevin Herkelman, Brad James, Pierre Lessard, Arnaud Samson, and Isabelle Cormier *Agriculture and Agri-Food Canada, Sherbrooke R & D Centre, Sherbrooke, QC J1M 1C8, Canada; † ‡ § Cooperative Research Farms, Richmond, VA 23235; Kalmbach Feeds, Upper Sandusky, OH 43351; Olymel, # $ St-Hyacinthe, QC J2S 4B6, Canada; Neovia, 02400 Château-Thierry, France; La COOP Fédérée, St-Romuald, QC G6W 5M6, Canada ABSTRACT: The impact of diet deprivation fol- (after standardization of litter size), and on days lowed by overallowance during gestation on met- 7, 14, and 21 (weaning). The TRT gilts gained less abolic status of pregnant gilts and their lactation BW than CTL gilts (17.3 vs. 31.7 kg; P < 0.01) performance was determined. Gilts were fed a from days 28 to 75 of gestation and more BW standard diet until day 27 of gestation and were (29.5 vs. 21.9 kg; P < 0.01) from days 75 to 110, subsequently reared under a control (CTL; n = 28) but their overall gain from mating to day 110 was or an experimental (treatment, TRT; n = 26) diet- lower (61.4 vs. 67.2 kg; P < 0.05). Metabolic status ary regimen. The experimental regimen provided during gestation was affected, with TRT gilts hav- 70% (restriction diet, RES) and 115% (overallow- ing less IGF-1 and urea, and more FFA than CTL ance diet, OVER) of the protein and NE contents gilts on day 75 (P < 0.01), and more urea on day provided by the CTL diet. The RES diet was given 110 (P < 0.01). Growth rate of suckling piglets, from days 28 to 74 of gestation followed by the sow lactation feed intake, and standard milk com- OVER diet from day 75 until farrowing. Blood position in late lactation (DM, fat, protein, lac- samples were obtained from all gilts on days 28, tose) were not affected by treatment (P > 0.10). In 75, and 110 of gestation, and on days 3 and 20 conclusion, diet deprivation of gilts as of day 28 of lactation to measure concentrations of IGF-1, of gestation followed by overfeeding from day 75 urea, FFA, and glucose. Milk samples were col- of gestation until farrowing did not improve lac- lected from 12 sows per treatment on day 19 of tation performance. It is likely that the compensa- lactation and sow feed intake was recorded daily tory growth that took place in late gestation was throughout lactation. Piglets were weighed at 24 h not adequate to illicit beneficial effects. Key words: diet deprivation, diet overallowance, gestation, milk yield, sows © Crown copyright 2018. Transl. Anim. Sci. 2018.2:162–168 doi: 10.1093/tas/txy012 INTRODUCTION Genetic selection and improved management The authors thank A.  Bernier and L.  Marier for their technologies have allowed sows to become more invaluable technical assistance, the staff at the CRF Sow productive in terms of litter size and milk yield. Research Farm for carrying out the project, and S.  Méthot However, milk yield needs to be further increased for statistical analyses. Sincere thanks are extended to La because growth rate of suckling piglets is not COOP Fédérée (St-Romuald, QC, Canada) and Cooperative optimal (Harrell et al., 1993). New feeding tech- Research Farms for financial and technical support. nologies in gestation, such as supplementation Corresponding author: chantal.farmer@agr.gc.ca with specific amino acids, phase-feeding, and use Received March 1, 2018. of high-fiber diets, have been used to improve Accepted April 10, 2018. Downloaded from https://academic.oup.com/tas/article-abstract/2/2/162/4969850 by Ed 'DeepDyve' Gillespie user on 19 June 2018 Gestation feeding and sow lactation 163 the performance of lactating sows (Sohn and diet was given from day 28 until day 74 of gesta- Maxwell, 1999). Since sow milk yield is affected tion, followed by the OVER diet from day 75 of by the number of mammary secretory cells pres- gestation until farrowing. These diets are described ent at the onset of lactation (Head and Williams, in Table 1. Representative feed samples were taken 1991), and rapid mammary accretion occurs in weekly from feed bins throughout the experiment the last third of gestation (Sorensen et al., 2002), for compositional analyses (Table  1). Gilts were several studies have focused on developing feed- ing strategies to stimulate mammary development Table 1. Composition of the experimental* and lac- in late gestation. Overfeeding a sow during gesta- tation (LACT) diets (as-fed) tion negatively affected mammary development Item CTL RES OVER LACT (Head and Williams, 1991), but diet deprivation Ingredient, g/kg followed by overallowance during the growing, Corn 486.2 38.8 584.0 579.1 finishing, and gestation phases increased sow milk Wheat, soft 100.0 119.9 100.0 - yield (Crenshaw et al., 1989). Recent results indi- Wheat middlings 180.0 250.0 22.3 100.0 cate that using a similar diet deprivation–over- Soybean meal (48% CP) 103.5 13.0 212.3 254.1 allowance dietary regimen in gestation does not Oat hulls 100.1 545.4 - - increase mammary development (Farmer et  al., Animal fat - - 50.0 23.1 2014). However, the level of diet deprivation was Ground limestone 14.9 14.0 14.1 15.6 too severe to bring about the expected compen- Dicalcium phosphate (21%) 5.6 6.5 6.9 14.7 satory growth, as evidenced by the lower BW of NaCl 6.3 6.1 6.3 5.0 Lys. HCl - 1.3 - 2.3 treated sows compared with that of control sows Met - 0.4 0.3 0.9 in late gestation (Farmer et  al., 2014). It was L-Thr - 0.6 0.4 0.9 hypothesized that reducing the duration of the L-Tryptophan - 0.1 - - diet deprivation imposed in early gestation while L-Valine - 0.5 - 0.7 using a similar overallowance in late gestation Choline (51.7%) 0.72 0.72 0.72 1.15 could stimulate mammary development via com- Trace mineral and vitamin premix 2.5 2.5 2.5 2.5 pensatory growth, hence increasing piglet growth Phytase 0.15 0.15 0.15 - rate in the subsequent lactation. Calculated composition DE, kcal/kg 3,115 2,286 3,624 3,432 NE, kcal/kg 2,232 1,559 2,566 2,525 MATERIALS AND METHODS CP, % 13.10 9.22 15.72 18.37 Fat, % 2.59 2.00 7.22 4.95 Animals were cared for according to a rec- Crude fiber, % 6.05 20.41 1.95 2.29 ommended code of practice (Agriculture and Total Lys, % 0.60 0.46 0.80 1.13 Agri-Food Canada, 1993) and procedures were Ca, % 0.75 0.75 0.75 0.95 approved by the Institutional Animal Care Committee P, % 0.57 0.54 0.50 0.75 of the Sherbrooke Research and Development Centre Met, % 0.21 0.17 0.27 0.37 of Agriculture and Agri-Food Canada. Met + Cys, % 0.46 0.35 0.54 0.67 Analyzed composition Animals and Treatments CP, % 12.87 9.95 16.63 18.73 Ca, % 0.76 0.71 0.77 0.96 Fifty-four Yorkshire × Landrace gilts were bred P, % 0.60 0.59 0.55 0.72 via AI using pools of semen from Duroc boars of Na, % 0.25 0.23 0.27 0.21 proven fertility. All gilts were fed 2.5 kg of a com- Mg, % 0.20 0.20 0.18 0.20 mercial diet containing 12% CP, 2232 kcal/kg NE, *Control (CTL), restriction (RES), and overallowance (OVER) 0.60% lysine, and 6.3% crude fiber from mating until diets fed from day 28 of gestation. The RES diet provided 70% and the day 27 of gestation. They were then fed 2.5 kg/d of OVER diet 115% of the CTL diet in the CP and NE contents. The RES diet was fed from days 28 until 74 of gestation followed by the OVER a control (CTL; n = 28) or an experimental (treat- diet for the remainder of gestation. ment, TRT; n = 26) dietary regimen. The CTL diet Provided the following per kilogram of diet: Cu (copper sulfate), was formulated to meet NRC requirements (2012). 15  mg; Zn (zinc sulfate), 125  mg; Se (sodium selenite), 0.3  mg; Mn The TRT regimen was designed to restrict growth (manganous sulfate), 40 mg; Fe (ferrous sulfate), 100 mg; I (ethylene diamine dihydroiodine), 0.5  mg; vitamin A, 10,000 IU; vitamin D, and then induce compensatory growth by provid- 1,400 IU; vitamin E, 60 IU; vitamin K, 3 mg; vitamin B , 20 µg; thia- ing 70% (restriction diet, RES) and 115% (overal- min, 1.5 mg; riboflavin, 6 mg; panthotenic acid, 25 mg; niacin, 30 mg; lowance diet, OVER), respectively, of the CP and folate, 8 mg; biotin, 0.5 mg; and pyridoxin, 3 mg. NE contents provided by the CTL diet. The RES Phyzyme XP (Danisco Animal Nutrition, Marlborough, UK). Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/2/2/162/4969850 by Ed 'DeepDyve' Gillespie user on 19 June 2018 164 Farmer et al. housed in individual gestation pens (0.61 × 1.90 m) fications as detailed previously ( Plante et al., 2011). and were weighed and backfat thickness was Validation for a plasma pool from sows was con- measured ultrasonically (WED-3000, Shenzhen ducted. Parallelism was 101.2% and average mass WELLD Medical Electronics Co., Ltd., Schenzhen, recovery was 101.3%. Sensitivity of the assay was China) at P2 of the last rib on days 28, 75, and 110 0.10 ng/mL. The intra-assay and interassay CV were of gestation, and on days 2 and 20 of lactation. 4.21% and 4.53%, respectively. Glucose was meas- Jugular blood samples were obtained from all gilts ured by an enzymatic colorimetric method with a on days 28, 75, and 110 of gestation and on days 3 commercial kit (Wako Pure Chemicals Industries and 20 of lactation. Ltd, Richmond, VA). Intra-assay and interas- Throughout lactation, sows were housed in a say CV were 2.00% and 1.25%, respectively. Urea 1.5  ×  2.1 m pen and were fed a commercial lac- was measured by colorimetric analysis using an tation diet (Table  1) in two equal meals at a rate automatic analyzer (Auto-Analyser 3; Technicon of 2.72  kg/d on the day of farrowing (day 0), Instruments Inc., Tarrytown, NY) according to 4.08 kg/d on day 1, 5.90 kg/d on day 2, 7.71 kg/d the method of Huntington (1984). Intra-assay on day 3, and then ad libitum for the remainder and interassay CV were 1.18% and 1.42%, respec- of lactation. Sow feed intake was recorded daily tively. Concentrations of FFA were also measured throughout lactation. Litter size was noted at by colorimetry with a commercial kit (Wako Pure birth and standardized to 12 or 13 piglets (within Chemicals industries Ltd). Intra-assay and interas- treatment group) within 24  h of birth. Piglets say CV were 3.05% and 6.79%, respectively. were weighed at 24 h (after standardization of lit- ter size), and on days 7, 14, and 21 (weaning) of Milk Composition lactation. Piglets had no access to dry feed while Whole milk was analyzed for DM, protein, suckling so that weight gain could provide an esti- fat, and lactose contents. DM was measured using mate of milk yield. Mortality rate was recorded forced air oven drying (method 925.23; AOAC, daily and reported for the first 48 h after farrowing 2005). Protein content was determined in dupli- and for the entire lactation period. Representative cates with the micro-Kjeldahl method (Kjeltec Auto milk samples were obtained from 12 CTL and 12 System; Tecator AB, Hoganas, Sweden) according TRT sows on day 19 of lactation, by collecting to AOAC Method 991.20 (AOAC, 2005), and fat milk from three functional glands (anterior, mid- was extracted using an established ether extraction dle, and posterior) encompassing both sides of method (method 905.02; AOAC, 2005). Lactose the udder after an intravenous injection of 1.0 mL was measured by a colorimetric method using a of oxytocin (20 IU/mL; P.V.U. Victoriaville, QC, commercial kit (Megazyme International Ireland Canada) was given. Piglets were separated from Ltd., Bray, Co. Wicklow, Ireland). Intra-assay and their dam for 45 min before oxytocin was injected. interassay CV were 2.09% and 1.17%, respectively. The post-weaning interval to estrus was recorded for all sows. Statistical Analyses Blood Handling and Assays The MIXED procedure of SAS (SAS Inst. Inc., Blood samples collected in gestation or lacta- Cary, NC) was used for statistical analyses. The tion were used to measure concentrations of IGF-1, univariate model used for sow backfat thickness urea, FFA, and glucose. Blood sampling was done and BW, milk composition, piglet BW, and IGF-1 between 0800 and 1000 h. Blood samples (30 mL) and metabolic variables in blood included the effect were collected in EDTA-tubes (Becton Dickinson of treatment, with the residual error being the and Cie, Rutherford, NJ), except those for glucose error term used to test main effects of treatment. analyses (6 mL), which were collected into tubes con- Repeated measures ANOVA with the factors treat- taining 12.0 mg of potassium oxalate and 15.0 mg ment (the error term being sow within treatment) of sodium fluoride to inhibit glycolysis. All sam - and sampling day (the residual error being the error ples were put on ice and centrifuged within 20 min term) and the treatment × day interaction were also at 1,800  × g for 12  min at 4  °C, and plasma was carried out on piglet BW and all blood variables. immediately recovered and frozen at −20  °C until The nonparametric Wilcoxon test was used to look assayed. Concentrations of IGF-1 were measured at treatment effect on litter size, and the Cochran– with a commercial kit for humans (ALPCO 26-G; Mantel–Haenszel test was used to determine treat- ALPCO Diagnostics, Salem, NH) with small modi- ment effects on the frequency of stillborn piglets Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/2/2/162/4969850 by Ed 'DeepDyve' Gillespie user on 19 June 2018 Gestation feeding and sow lactation 165 and piglet mortality during the 48 h postpartum or gilts weighed 12.2 kg less than CTL gilts (Table  2, the whole lactation period. Data in text and tables P  <  0.01), but backfat thickness was not different are presented as least squares means ± SEM unless between gilts from both groups (Table 2, P > 0.10). specified otherwise. During the period of feed overallowance (days 75 to 110 of gestation), TRT gilts gained more BW than CTL gilts (Table 2, P < 0.01) and lost less backfat RESULTS (Table 2, P < 0.01). On day 110 of gestation, TRT During the period when the RES diet was fed and CTL gilts did not differ in BW and backfat (days 28 to 74 of gestation), TRT gilts gained less (Table  2, P > 0.10). The BW gain over the whole BW than CTL gilts (Table  2, P  <  0.01) and also period from days 28 until 110 of gestation was lost backfat, whereas CTL gilts gained backfat lower for TRT than CTL gilts (Table 2, P < 0.05), (Table  2, P  <  0.01). On day 75 of gestation, TRT whereas the change in backfat thickness did not dif- fer across treatments (Table 2, P > 0.10). There was a significant treatment × day inter - Table  2. Weight, backfat thickness and lactation action for circulating concentrations of urea, FFA, feed intakes of sows [28 control (CTL) and 26 and IGF-1 over gestation (Table  3, P  <  0.01). treated (TRT) sows]* Analyses done per sampling day showed that at the Groups † Table  3. Circulating concentrations of urea, FFA, Item CTL TRT SEM glucose, and IGF-1 in pregnant and lactating prim- BW, kg iparous sows [28 control (CTL) and 27 treated Mating 152.3 153.5 2.9 Day 28 of gestation 165.9 168.0 2.8 (TRT) sows]* a b Day 75 of gestation 197.6 185.4 3.1 Groups Day 110 of gestation 219.5 214.9 3.3 a b Item CTL TRT SEM Gain, days 28 to 75 31.7 17.3 1.2 a b Urea, mmol/L Gain, days 75 to 110 21.9 29.5 1.0 a b Day 28 of gestation 5.01 4.94 0.18 Gain, days 28 to 110 53.6 46.9 1.4 a b c d Day 75 of gestation 4.80 4.03 0.19 Gain, mating to day 110 67.2 61.4 1.8 a b Day 110 of gestation 4.76 5.66 0.27 Day 2 of lactation 200.3 197.4 2.6 Day 3 of lactation 6.11 6.98 0.49 Day 20 of lactation 187.0 186.2 3.3 Day 20 of lactation 9.95 10.69 0.52 Loss from days 2 to 20 14.4 11.3 1.8 FFA, µEq/L Backfat, mm Day 28 of gestation 137.34 101.17 18.78 Mating 16.6 17.0 0.8 a b Day 75 of gestation 125.27 275.20 33.14 Day 28 of gestation 17.9 18.3 0.8 Day 110 of gestation 342.02 282.55 38.90 Day 75 of gestation 18.5 16.9 0.8 Day 3 of lactation 205.87 252.19 54.55 Day 110 of gestation 17.1 17.0 0.8 c d a b Day 20 of lactation 285.27 171.53 54.99 Difference, days 28 to 75 0.6 -1.4 0.4 a b Glucose, mmol/L Difference, days 75 to 110 -1.8 -0.3 0.4 Day 28 of gestation 3.22 3.33 0.07 Difference, days 28 to 110 -0.8 -1.3 0.5 Day 75 of gestation 3.25 3.27 0.06 Difference, mating to day 110 0.5 -0.1 0.5 Day 110 of gestation 3.32 3.31 0.85 Day 2 of lactation 16.7 16.7 0.8 Day 3 of lactation 4.15 3.91 0.11 Day 20 of lactation 13.7 14.3 0.7 c d Day 20 of lactation 3.76 4.05 0.12 Difference, days 2 to 20 -3.5 -2.7 0.5 IGF-1, ng/mL Average daily feed intake, kg Day 28 of gestation 82.6 84.2 2.9 Week 1 of lactation 4.38 4.47 0.24 a b Day 75 of gestation 52.2 37.4 2.5 Week 2 of lactation 5.74 5.85 0.25 Day 110 of gestation 42.2 44.4 2.4 Week 3 of lactation 5.61 6.27 0.37 Day 3 of lactation 102.9 97.9 6.7 Total lactation feed intake, kg 105.15 109.37 5.33 Day 20 of lactation 101.5 102.1 9.3 a,b Means within a row with different superscripts differ (P < 0.01). a,b c,d Means within a row with different superscripts differ (P < 0.01). Means within a row with different superscripts differ (P < 0.05). c,d Means within a row with different superscripts tend to differ *CTL  =  control, and TRT  =  treatment regimen, which provided (P < 0.10). 70% (restriction) and 115% (overallowance) of the CTL diet in the CP and NE contents. The restriction diet was fed from days 28 until 74 *CTL = control, and TRT = treatment regimen, which provided 70 of gestation followed by the overallowance diet for the remainder of (restriction) and 115% (overallowance) of the CTL diet in the CP and gestation. Sows were fed 2.5 kg/d throughout gestation and there were NE contents. The restriction diet was fed from days 28 until 74 of gesta- no refusals. tion followed by the overallowance diet for the remainder of gestation. † † Maximum value. Maximum value. Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/2/2/162/4969850 by Ed 'DeepDyve' Gillespie user on 19 June 2018 166 Farmer et al. end of feeding the RES diet (day 75 of gestation), and protein intake from days 28 until 74 of gestation TRT gilts had less urea, more than double the con- and then overfeeding gilts from days 75 until 110 of centrations of FFA and less IGF-1 than CTL gilts gestation for subsequent lactation performance. An (Table  3, P  <  0.01). After feeding the OVER diet earlier trial suggested that diet deprivation followed (day 110), TRT gilts had more urea than CTL gilts by overallowance during the growing, finishing, (Table  3, P  <  0.01). During lactation, there was a and gestation phases could be beneficial in terms of significant treatment × day interaction for glu - milk yield and mammary gene expression in swine cose (P  <  0.01), a tendency for a treatment × day (Crenshaw et  al., 1989). Yet, a recent experiment interaction for FFA (P < 0.10), and a day effect for showed that when a similar diet deprivation-over- urea (P < 0.01), with values increasing as lactation allowance regimen as described by Crenshaw et al. advanced (Table  3). There were no changes in any (1989) was used in gestation only, there was no measured variables due to treatment on day 3 of increase in mammary development of late-pregnant lactation (P > 0.10), but on day 20 of lactation, gilts (Farmer et al., 2014). It was hypothesized that TRT sows tended to have less FFA and more glu- the level of diet deprivation imposed in early ges- cose than CTL sows (Table 3, P < 0.10). There was tation was too severe for the compensatory growth no change in weekly or total lactation feed intake in late gestation to have any beneficial effects. This of sows across treatments (Table  2; P > 0.10) and was suggested by the 12.9 kg lower BW of treated milk DM, fat, protein, and lactose contents on gilts compared with control gilts on day 110 of ges- day 19 of lactation were not altered by treatment tation (Farmer et al., 2014). In the current trial, the (Table 4; P > 0.10). duration of the diet deprivation was reduced, start- The number of live-born piglets did not vary ing on day 28 of gestation, while an overallowance between treatments, being 13.7 and 12.5  ±  0.5 for similar to previous trials was used in later gestation. CTL and TRT sows, respectively (P  =  0.1). Piglet As expected, concentrations of FFA at the end of BW at birth or on days 1, 7, 14, or 21 of lactation the restricted period were increased due to greater was not affected by treatment, nor was BW gain use of energy reserves (Barb et  al., 1997; Farmer over the lactation period altered (Table 5; P > 0.10). et al., 2014; Ren et al., 2017) and urea was reduced There was no difference between treatments on the indicative of a lower supply of proteins (Pedersen frequency of piglet mortality that occurred either in et al., 2016). The 4.6 kg difference in BW between the first 48 h postpartum (12 vs. 11 sows with dead TRT and CTL gilts on day 110 of gestation in the piglets for CTL and TRT, respectively; P > 0.10) current experiment was much lower than that pre- or over the whole lactation period (7 vs. 10 sows viously seen (Farmer et al., 2014) and was not sig- with dead piglets for CTL and TRT, respectively; nificant. Nevertheless, the increase in BW of gilts P > 0.10), but the incidence of stillborn was greater from days 75 to 110 of gestation was not as large as in CTL than TRT sows (13 vs. 7 sows with stillborn the reduction in BW from days 28 until 75, so that piglets for CTL and TRT, respectively; P < 0.05). even though compensatory growth did take place in DISCUSSION Table 5. Weight of piglets* Groups Findings from the current experiment indicate that there is no beneficial effect of restricting energy Item CTL TRT SEM BW, kg Table  4. Milk composition on day 19 of lactation Birth 1.43 (12.9) 1.50 (12.7) 0.05 (n = 12)* Day 1 of lactation 1.45 (12.3) 1.51 (12.2) 0.05 Day 7 of lactation 2.75 (12.3) 2.80 (11.7) 0.10 Groups Day 14 of lactation 4.52 (11.9) 4.51 (11.7) 0.13 Item CTL TRT SEM Day 21 of lactation 6.10 (11.9) 6.02 (11.7) 0.20 DM, % 18.78 19.52 0.42 BW gain, kg Fat, % 7.30 7.78 0.37 Days 1 to 21 4.65 4.50 0.17 Protein, % 5.38 5.65 0.18 Average numbers of piglets per litter are shown in parenthesis. Lactose, % 4.86 4.71 0.11 *CTL  =  control, and TRT  =  treatment regimen, which provided 70 (restriction) and 115% (overallowance) of the CTL diet in the CP *CTL = control, and TRT = treatment regimen, which provided 70 and NE contents. The restriction diet was fed from days 28 until 74 (restriction) and 115% (overallowance) of the CTL diet in the CP and of gestation followed by the overallowance diet for the remainder of NE contents. The restriction diet was fed from days 28 until 74 of gesta- gestation. tion followed by the overallowance diet for the remainder of gestation. † † Maximum value. Maximum value. Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/2/2/162/4969850 by Ed 'DeepDyve' Gillespie user on 19 June 2018 Gestation feeding and sow lactation 167 late gestation it was likely not important enough to across treatments are in agreement with the unal- stimulate mammary development. This hypothesis tered milk composition. The similar birthweight of is substantiated by the fact that circulating IGF-1 piglets across treatments also indicates that fetal concentrations were lower in TRT than CTL gilts growth was not affected even though there may on day 75 of gestation and were not greater on day have been a positive effect on the incidence of still- 110 of gestation. Lower IGF-1 concentrations in borns. Yet, the number of animals used was too animals with a slower BW gain were expected due low to determine such an effect and this needs to to the positive relationship between IGF-1 concen- be corroborated in future studies. Ren et al. (2017) trations and growth rate in growing pigs (Owens recently demonstrated that feed restriction or over- et al., 1999). Taking into account the important role feeding of gilts during three short periods of gesta- of IGF-1 for mammary development in rodents tion did not affect litter performance. Even though (Kleinberg and Barcellos-Hoff, 2011) and for mito- growth performance of gilts was increased during genesis of mammary epithelial cells in ruminants those periods when they were fed twice the main- (Forsyth, 1996), one might suggest that for a late tenance requirements, it did not lead to beneficial gestation compensatory feeding strategy to have effects during the subsequent lactation. Therefore, beneficial effects on lactation performance it would altering growth rate of gilts during gestation may need to increase IGF-1 concentrations. not be an adequate strategy to enhance mammary The compensatory feeding regimen was imposed development or the changes incurred need to be in gestation only in the present trial, because when more prominent to have an effect. a feeding regimen similar to that of Crenshaw et al. In conclusion, diet deprivation of gilts as of (1989) was used in the growing-finishing period, day 28 of gestation followed by overfeeding from there was no beneficial effect on mammary devel - day 75 of gestation until farrowing did not improve opment either at puberty (Farmer et al., 2012a) or lactation performance. It is likely that the compen- at 110 d of gestation (Lyvers-Peffer and Rozeboom, satory growth that took place in late gestation was 2001; Farmer et  al., 2012b). In studies by Farmer not adequate to illicit beneficial effects on mam - et al. (2012a, 2012b), the greater BW gain observed mary development because IGF-1 concentrations by Crenshaw (1990) in treated gilts at the end of were not increased. the growing-finishing period was not seen and in the study by Lyvers-Peffer and Rozeboom (2001), LITERATURE CITED BW of gilts was not reported. In fact, gilts sub- Agriculture and Agri-Food Canada. 1993. Recommended code jected to the restricted and overallowance feeding of practice for the care and handling of farm animals - regime weighed 6.2 kg less at puberty than control Pigs. Publ. No. 1898E. Ottawa, ON (Canada): Agriculture and Agri-Food Canada. gilts (Farmer et al., 2012a). Differences in the type AOAC. 2005. Official methods of analysis international. 18th of fiber used and in feed composition were thought ed. Arlington (VA): Assoc. Off. Anal. Chem. to be the cause of this discrepancy. Previous results Barb, C. R., R. R. Kraeling, G. B. Rampacek, and C. R. Dove. (Farmer et  al. 2012a, 2012b) combined with cur- 1997. Metabolic changes during the transition from the rent results therefore question the hypothesis that fed to the acute feed-deprived state in prepuberal and the beneficial effect seen by Crenshaw et al. (1989) mature gilts. J. Anim. Sci. 75:781–789. Crenshaw, J. D. 1990. Feeding gilts to enhance lactational per- on sow milk yield was mostly due to compensa- formance. Fargo (ND): North Dakota State Univ. Swine tory growth in late gestation. It may be that a feed- Res. Rep. Agric. Exp. Stn. Ext. Services. p.12–23. ing regime that could bring about the 8.2  kg BW Crenshaw, J. D., C. S. Park, P. M. Swantek, W. L. Keller, and R. advantage of treated gilts at the end of the grow- C. Zimprich. 1989. Lactation response of gilts to a phased ing-finishing period in Crenshaw’s study would be feeding regimen designed to induce compensatory growth. J. Anim. Sci. 67(Suppl. 2):107–108. (Abstr.) beneficial for mammary development and future Farmer, C., M. F. Palin, and Y. Martel-Kennes. 2012a. Impact milk yield. However, circulating concentrations of of diet deprivation and subsequent over-allowance during IGF-1 were not measured in the study by Crenshaw prepuberty. Part 1. Effects on growth performance, metab- et al. (1989), so it is unknown if the beneficial effect olite status, and mammary gland development in gilts. J. on sow milk yield was mediated through this growth Anim. Sci. 90:863–871. doi:10.2527/jas.2011-4131 factor. Farmer, C., M. F. Palin, and Y. Martel-Kennes. 2012b. Impact of diet deprivation and subsequent over-allowance during The lack of treatment effect on circulating prepuberty. Part 2.  Effects on mammary gland develop- concentrations of IGF-1 on day 110 of gestation, ment and lactation performance of sows. J. Anim. Sci. the presence of only tendencies for decreased FFA 90:872–880. doi:10.2527/jas.2011-4480 and increased glucose in TRT vs. CTL gilts on that Farmer, C., M. F. Palin, and Y. Martel-Kennes. 2014. Impact same day, and the similar lactation feed intakes of diet deprivation and subsequent overallowance during Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/2/2/162/4969850 by Ed 'DeepDyve' Gillespie user on 19 June 2018 168 Farmer et al. gestation on mammary gland development and lacta- National Research Council. 2012. Composition of feed tion performance. J. Anim. Sci. 92:141–151. doi:10.2527/ ingredients. Nutrient requirements of swine. 11th ed. jas.2013-6558 Washington, DC: National Academy Press. pp 239–367. Forsyth, I. A. 1996. The insulin-like growth factor and epi- Owens, P. C., K. L.  Gatford, P. E.  Walton, W.  Morley, and dermal growth factor families in mammary cell growth in R. G. Campbell. 1999. The relationship between endogen- ruminants: action and interaction with hormones. J. Dairy ous insulin-like growth factors and growth in pigs. J. Sci. 79:1085–1096. doi:10.3168/jds.S0022-0302(96)76462-7 Anim. Sci. 77:2098–2103. Harrell, R. J., M. J. Thomas, and R. D. Boyd. 1993. Limitations Pedersen, T. F., T. S.  Bruun, T.  Feyera, U. K.  Larsen, and of sow milk yield on baby pig growth. Proc. 1993 Cornell P. K.Theil. 2016. A two-diet feeding regime for lactating Nutr. Conf. for Feed Manufacturers p. 156–164. sows reduced nutrient deficiency in early lactation and Head, R. H., and I. H. Williams. 1991. Mammogenesis is influ - improved milk yield. Livest. Sci. 191:165–173. enced by pregnancy nutrition. In: Manipulating pig pro- Plante, P-A., J-P. Laforest, and C. Farmer. 2011. Effect of sup- duction III. VIC, Australia: Australasian Pig Sci. Assoc., plementing the diet of lactating sows with NuPro® on Werribee. p. 33. their performances and that of their piglets. Can. J. Anim. Huntington, G. B. 1984. Net absorption of glucose and nitrog- Sci. 91:295–300. enous compounds by lactating Holstein cows. J. Dairy Sci. Ren, P., X.  J. Yang, S. Q. Cui, J. S. Kim, D. Menon, and 67:1919–1927. doi:10.3168/jds.S0022-0302(84)81525-8 S. K. Baidoo. 2017. Effects of different feeding levels dur- Kleinberg, D. L., and M. H.  Barcellos-Hoff. 2011. The piv- ing three short periods of gestation on gilt and litter per- otal role of insulin-like growth factor I  in normal mam- formance, nutrient digestibility, and energy homeostasis in mary development. Endocrinol. Metab. Clin. N.  Amer. gilts. J. Anim. Sci. 95:1232–1242. doi:10.2527/jas.2016.1208 40:461–471. Sohn, K. S., and C. V.  Maxwell. 1999. New technologies for Lyvers-Peffer, P. A., and D. W. Rozeboom. 2001. The effects of sow nutrition and management – Review. Asian-Aus. a growth-altering pre-pubertal feeding regimen on mam- J. Anim. Sci. 12:956–965. mary development and parity-one lactation potential in Sorensen, M. T., K.  Sejrsen, and S.  Purup. 2002. Mammary swine. Livest. Prod. Sci. 70:167–173. gland development in gilts. Livest. Prod. Sci. 75:143–148. 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Impact of diet deprivation and subsequent overallowance during gestation on lactation performance of primiparous sows

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Impact of diet deprivation and subsequent overallowance during gestation on lactation performance of primiparous sows ,2 † ‡ § # Chantal Farmer,* Kevin Herkelman, Brad James, Pierre Lessard, Arnaud Samson, and Isabelle Cormier *Agriculture and Agri-Food Canada, Sherbrooke R & D Centre, Sherbrooke, QC J1M 1C8, Canada; † ‡ § Cooperative Research Farms, Richmond, VA 23235; Kalmbach Feeds, Upper Sandusky, OH 43351; Olymel, # $ St-Hyacinthe, QC J2S 4B6, Canada; Neovia, 02400 Château-Thierry, France; La COOP Fédérée, St-Romuald, QC G6W 5M6, Canada ABSTRACT: The impact of diet deprivation fol- (after standardization of litter size), and on days lowed by overallowance during gestation on met- 7, 14, and 21 (weaning). The TRT gilts gained less abolic status of pregnant gilts and their lactation BW than CTL gilts (17.3 vs. 31.7 kg; P < 0.01) performance was determined. Gilts were fed a from days 28 to 75 of gestation and more BW standard diet until day 27 of gestation and were (29.5 vs. 21.9 kg; P < 0.01) from days 75 to 110, subsequently reared under a control (CTL; n = 28) but their overall gain from mating to day 110 was or an experimental (treatment, TRT; n = 26) diet- lower (61.4 vs. 67.2 kg; P < 0.05). Metabolic status ary regimen. The experimental regimen provided during gestation was affected, with TRT gilts hav- 70% (restriction diet, RES) and 115% (overallow- ing less IGF-1 and urea, and more FFA than CTL ance diet, OVER) of the protein and NE contents gilts on day 75 (P < 0.01), and more urea on day provided by the CTL diet. The RES diet was given 110 (P < 0.01). Growth rate of suckling piglets, from days 28 to 74 of gestation followed by the sow lactation feed intake, and standard milk com- OVER diet from day 75 until farrowing. Blood position in late lactation (DM, fat, protein, lac- samples were obtained from all gilts on days 28, tose) were not affected by treatment (P > 0.10). In 75, and 110 of gestation, and on days 3 and 20 conclusion, diet deprivation of gilts as of day 28 of lactation to measure concentrations of IGF-1, of gestation followed by overfeeding from day 75 urea, FFA, and glucose. Milk samples were col- of gestation until farrowing did not improve lac- lected from 12 sows per treatment on day 19 of tation performance. It is likely that the compensa- lactation and sow feed intake was recorded daily tory growth that took place in late gestation was throughout lactation. Piglets were weighed at 24 h not adequate to illicit beneficial effects. Key words: diet deprivation, diet overallowance, gestation, milk yield, sows © Crown copyright 2018. Transl. Anim. Sci. 2018.2:162–168 doi: 10.1093/tas/txy012 INTRODUCTION Genetic selection and improved management The authors thank A.  Bernier and L.  Marier for their technologies have allowed sows to become more invaluable technical assistance, the staff at the CRF Sow productive in terms of litter size and milk yield. Research Farm for carrying out the project, and S.  Méthot However, milk yield needs to be further increased for statistical analyses. Sincere thanks are extended to La because growth rate of suckling piglets is not COOP Fédérée (St-Romuald, QC, Canada) and Cooperative optimal (Harrell et al., 1993). New feeding tech- Research Farms for financial and technical support. nologies in gestation, such as supplementation Corresponding author: chantal.farmer@agr.gc.ca with specific amino acids, phase-feeding, and use Received March 1, 2018. of high-fiber diets, have been used to improve Accepted April 10, 2018. Downloaded from https://academic.oup.com/tas/article-abstract/2/2/162/4969850 by Ed 'DeepDyve' Gillespie user on 19 June 2018 Gestation feeding and sow lactation 163 the performance of lactating sows (Sohn and diet was given from day 28 until day 74 of gesta- Maxwell, 1999). Since sow milk yield is affected tion, followed by the OVER diet from day 75 of by the number of mammary secretory cells pres- gestation until farrowing. These diets are described ent at the onset of lactation (Head and Williams, in Table 1. Representative feed samples were taken 1991), and rapid mammary accretion occurs in weekly from feed bins throughout the experiment the last third of gestation (Sorensen et al., 2002), for compositional analyses (Table  1). Gilts were several studies have focused on developing feed- ing strategies to stimulate mammary development Table 1. Composition of the experimental* and lac- in late gestation. Overfeeding a sow during gesta- tation (LACT) diets (as-fed) tion negatively affected mammary development Item CTL RES OVER LACT (Head and Williams, 1991), but diet deprivation Ingredient, g/kg followed by overallowance during the growing, Corn 486.2 38.8 584.0 579.1 finishing, and gestation phases increased sow milk Wheat, soft 100.0 119.9 100.0 - yield (Crenshaw et al., 1989). Recent results indi- Wheat middlings 180.0 250.0 22.3 100.0 cate that using a similar diet deprivation–over- Soybean meal (48% CP) 103.5 13.0 212.3 254.1 allowance dietary regimen in gestation does not Oat hulls 100.1 545.4 - - increase mammary development (Farmer et  al., Animal fat - - 50.0 23.1 2014). However, the level of diet deprivation was Ground limestone 14.9 14.0 14.1 15.6 too severe to bring about the expected compen- Dicalcium phosphate (21%) 5.6 6.5 6.9 14.7 satory growth, as evidenced by the lower BW of NaCl 6.3 6.1 6.3 5.0 Lys. HCl - 1.3 - 2.3 treated sows compared with that of control sows Met - 0.4 0.3 0.9 in late gestation (Farmer et  al., 2014). It was L-Thr - 0.6 0.4 0.9 hypothesized that reducing the duration of the L-Tryptophan - 0.1 - - diet deprivation imposed in early gestation while L-Valine - 0.5 - 0.7 using a similar overallowance in late gestation Choline (51.7%) 0.72 0.72 0.72 1.15 could stimulate mammary development via com- Trace mineral and vitamin premix 2.5 2.5 2.5 2.5 pensatory growth, hence increasing piglet growth Phytase 0.15 0.15 0.15 - rate in the subsequent lactation. Calculated composition DE, kcal/kg 3,115 2,286 3,624 3,432 NE, kcal/kg 2,232 1,559 2,566 2,525 MATERIALS AND METHODS CP, % 13.10 9.22 15.72 18.37 Fat, % 2.59 2.00 7.22 4.95 Animals were cared for according to a rec- Crude fiber, % 6.05 20.41 1.95 2.29 ommended code of practice (Agriculture and Total Lys, % 0.60 0.46 0.80 1.13 Agri-Food Canada, 1993) and procedures were Ca, % 0.75 0.75 0.75 0.95 approved by the Institutional Animal Care Committee P, % 0.57 0.54 0.50 0.75 of the Sherbrooke Research and Development Centre Met, % 0.21 0.17 0.27 0.37 of Agriculture and Agri-Food Canada. Met + Cys, % 0.46 0.35 0.54 0.67 Analyzed composition Animals and Treatments CP, % 12.87 9.95 16.63 18.73 Ca, % 0.76 0.71 0.77 0.96 Fifty-four Yorkshire × Landrace gilts were bred P, % 0.60 0.59 0.55 0.72 via AI using pools of semen from Duroc boars of Na, % 0.25 0.23 0.27 0.21 proven fertility. All gilts were fed 2.5 kg of a com- Mg, % 0.20 0.20 0.18 0.20 mercial diet containing 12% CP, 2232 kcal/kg NE, *Control (CTL), restriction (RES), and overallowance (OVER) 0.60% lysine, and 6.3% crude fiber from mating until diets fed from day 28 of gestation. The RES diet provided 70% and the day 27 of gestation. They were then fed 2.5 kg/d of OVER diet 115% of the CTL diet in the CP and NE contents. The RES diet was fed from days 28 until 74 of gestation followed by the OVER a control (CTL; n = 28) or an experimental (treat- diet for the remainder of gestation. ment, TRT; n = 26) dietary regimen. The CTL diet Provided the following per kilogram of diet: Cu (copper sulfate), was formulated to meet NRC requirements (2012). 15  mg; Zn (zinc sulfate), 125  mg; Se (sodium selenite), 0.3  mg; Mn The TRT regimen was designed to restrict growth (manganous sulfate), 40 mg; Fe (ferrous sulfate), 100 mg; I (ethylene diamine dihydroiodine), 0.5  mg; vitamin A, 10,000 IU; vitamin D, and then induce compensatory growth by provid- 1,400 IU; vitamin E, 60 IU; vitamin K, 3 mg; vitamin B , 20 µg; thia- ing 70% (restriction diet, RES) and 115% (overal- min, 1.5 mg; riboflavin, 6 mg; panthotenic acid, 25 mg; niacin, 30 mg; lowance diet, OVER), respectively, of the CP and folate, 8 mg; biotin, 0.5 mg; and pyridoxin, 3 mg. NE contents provided by the CTL diet. The RES Phyzyme XP (Danisco Animal Nutrition, Marlborough, UK). Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/2/2/162/4969850 by Ed 'DeepDyve' Gillespie user on 19 June 2018 164 Farmer et al. housed in individual gestation pens (0.61 × 1.90 m) fications as detailed previously ( Plante et al., 2011). and were weighed and backfat thickness was Validation for a plasma pool from sows was con- measured ultrasonically (WED-3000, Shenzhen ducted. Parallelism was 101.2% and average mass WELLD Medical Electronics Co., Ltd., Schenzhen, recovery was 101.3%. Sensitivity of the assay was China) at P2 of the last rib on days 28, 75, and 110 0.10 ng/mL. The intra-assay and interassay CV were of gestation, and on days 2 and 20 of lactation. 4.21% and 4.53%, respectively. Glucose was meas- Jugular blood samples were obtained from all gilts ured by an enzymatic colorimetric method with a on days 28, 75, and 110 of gestation and on days 3 commercial kit (Wako Pure Chemicals Industries and 20 of lactation. Ltd, Richmond, VA). Intra-assay and interas- Throughout lactation, sows were housed in a say CV were 2.00% and 1.25%, respectively. Urea 1.5  ×  2.1 m pen and were fed a commercial lac- was measured by colorimetric analysis using an tation diet (Table  1) in two equal meals at a rate automatic analyzer (Auto-Analyser 3; Technicon of 2.72  kg/d on the day of farrowing (day 0), Instruments Inc., Tarrytown, NY) according to 4.08 kg/d on day 1, 5.90 kg/d on day 2, 7.71 kg/d the method of Huntington (1984). Intra-assay on day 3, and then ad libitum for the remainder and interassay CV were 1.18% and 1.42%, respec- of lactation. Sow feed intake was recorded daily tively. Concentrations of FFA were also measured throughout lactation. Litter size was noted at by colorimetry with a commercial kit (Wako Pure birth and standardized to 12 or 13 piglets (within Chemicals industries Ltd). Intra-assay and interas- treatment group) within 24  h of birth. Piglets say CV were 3.05% and 6.79%, respectively. were weighed at 24 h (after standardization of lit- ter size), and on days 7, 14, and 21 (weaning) of Milk Composition lactation. Piglets had no access to dry feed while Whole milk was analyzed for DM, protein, suckling so that weight gain could provide an esti- fat, and lactose contents. DM was measured using mate of milk yield. Mortality rate was recorded forced air oven drying (method 925.23; AOAC, daily and reported for the first 48 h after farrowing 2005). Protein content was determined in dupli- and for the entire lactation period. Representative cates with the micro-Kjeldahl method (Kjeltec Auto milk samples were obtained from 12 CTL and 12 System; Tecator AB, Hoganas, Sweden) according TRT sows on day 19 of lactation, by collecting to AOAC Method 991.20 (AOAC, 2005), and fat milk from three functional glands (anterior, mid- was extracted using an established ether extraction dle, and posterior) encompassing both sides of method (method 905.02; AOAC, 2005). Lactose the udder after an intravenous injection of 1.0 mL was measured by a colorimetric method using a of oxytocin (20 IU/mL; P.V.U. Victoriaville, QC, commercial kit (Megazyme International Ireland Canada) was given. Piglets were separated from Ltd., Bray, Co. Wicklow, Ireland). Intra-assay and their dam for 45 min before oxytocin was injected. interassay CV were 2.09% and 1.17%, respectively. The post-weaning interval to estrus was recorded for all sows. Statistical Analyses Blood Handling and Assays The MIXED procedure of SAS (SAS Inst. Inc., Blood samples collected in gestation or lacta- Cary, NC) was used for statistical analyses. The tion were used to measure concentrations of IGF-1, univariate model used for sow backfat thickness urea, FFA, and glucose. Blood sampling was done and BW, milk composition, piglet BW, and IGF-1 between 0800 and 1000 h. Blood samples (30 mL) and metabolic variables in blood included the effect were collected in EDTA-tubes (Becton Dickinson of treatment, with the residual error being the and Cie, Rutherford, NJ), except those for glucose error term used to test main effects of treatment. analyses (6 mL), which were collected into tubes con- Repeated measures ANOVA with the factors treat- taining 12.0 mg of potassium oxalate and 15.0 mg ment (the error term being sow within treatment) of sodium fluoride to inhibit glycolysis. All sam - and sampling day (the residual error being the error ples were put on ice and centrifuged within 20 min term) and the treatment × day interaction were also at 1,800  × g for 12  min at 4  °C, and plasma was carried out on piglet BW and all blood variables. immediately recovered and frozen at −20  °C until The nonparametric Wilcoxon test was used to look assayed. Concentrations of IGF-1 were measured at treatment effect on litter size, and the Cochran– with a commercial kit for humans (ALPCO 26-G; Mantel–Haenszel test was used to determine treat- ALPCO Diagnostics, Salem, NH) with small modi- ment effects on the frequency of stillborn piglets Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/2/2/162/4969850 by Ed 'DeepDyve' Gillespie user on 19 June 2018 Gestation feeding and sow lactation 165 and piglet mortality during the 48 h postpartum or gilts weighed 12.2 kg less than CTL gilts (Table  2, the whole lactation period. Data in text and tables P  <  0.01), but backfat thickness was not different are presented as least squares means ± SEM unless between gilts from both groups (Table 2, P > 0.10). specified otherwise. During the period of feed overallowance (days 75 to 110 of gestation), TRT gilts gained more BW than CTL gilts (Table 2, P < 0.01) and lost less backfat RESULTS (Table 2, P < 0.01). On day 110 of gestation, TRT During the period when the RES diet was fed and CTL gilts did not differ in BW and backfat (days 28 to 74 of gestation), TRT gilts gained less (Table  2, P > 0.10). The BW gain over the whole BW than CTL gilts (Table  2, P  <  0.01) and also period from days 28 until 110 of gestation was lost backfat, whereas CTL gilts gained backfat lower for TRT than CTL gilts (Table 2, P < 0.05), (Table  2, P  <  0.01). On day 75 of gestation, TRT whereas the change in backfat thickness did not dif- fer across treatments (Table 2, P > 0.10). There was a significant treatment × day inter - Table  2. Weight, backfat thickness and lactation action for circulating concentrations of urea, FFA, feed intakes of sows [28 control (CTL) and 26 and IGF-1 over gestation (Table  3, P  <  0.01). treated (TRT) sows]* Analyses done per sampling day showed that at the Groups † Table  3. Circulating concentrations of urea, FFA, Item CTL TRT SEM glucose, and IGF-1 in pregnant and lactating prim- BW, kg iparous sows [28 control (CTL) and 27 treated Mating 152.3 153.5 2.9 Day 28 of gestation 165.9 168.0 2.8 (TRT) sows]* a b Day 75 of gestation 197.6 185.4 3.1 Groups Day 110 of gestation 219.5 214.9 3.3 a b Item CTL TRT SEM Gain, days 28 to 75 31.7 17.3 1.2 a b Urea, mmol/L Gain, days 75 to 110 21.9 29.5 1.0 a b Day 28 of gestation 5.01 4.94 0.18 Gain, days 28 to 110 53.6 46.9 1.4 a b c d Day 75 of gestation 4.80 4.03 0.19 Gain, mating to day 110 67.2 61.4 1.8 a b Day 110 of gestation 4.76 5.66 0.27 Day 2 of lactation 200.3 197.4 2.6 Day 3 of lactation 6.11 6.98 0.49 Day 20 of lactation 187.0 186.2 3.3 Day 20 of lactation 9.95 10.69 0.52 Loss from days 2 to 20 14.4 11.3 1.8 FFA, µEq/L Backfat, mm Day 28 of gestation 137.34 101.17 18.78 Mating 16.6 17.0 0.8 a b Day 75 of gestation 125.27 275.20 33.14 Day 28 of gestation 17.9 18.3 0.8 Day 110 of gestation 342.02 282.55 38.90 Day 75 of gestation 18.5 16.9 0.8 Day 3 of lactation 205.87 252.19 54.55 Day 110 of gestation 17.1 17.0 0.8 c d a b Day 20 of lactation 285.27 171.53 54.99 Difference, days 28 to 75 0.6 -1.4 0.4 a b Glucose, mmol/L Difference, days 75 to 110 -1.8 -0.3 0.4 Day 28 of gestation 3.22 3.33 0.07 Difference, days 28 to 110 -0.8 -1.3 0.5 Day 75 of gestation 3.25 3.27 0.06 Difference, mating to day 110 0.5 -0.1 0.5 Day 110 of gestation 3.32 3.31 0.85 Day 2 of lactation 16.7 16.7 0.8 Day 3 of lactation 4.15 3.91 0.11 Day 20 of lactation 13.7 14.3 0.7 c d Day 20 of lactation 3.76 4.05 0.12 Difference, days 2 to 20 -3.5 -2.7 0.5 IGF-1, ng/mL Average daily feed intake, kg Day 28 of gestation 82.6 84.2 2.9 Week 1 of lactation 4.38 4.47 0.24 a b Day 75 of gestation 52.2 37.4 2.5 Week 2 of lactation 5.74 5.85 0.25 Day 110 of gestation 42.2 44.4 2.4 Week 3 of lactation 5.61 6.27 0.37 Day 3 of lactation 102.9 97.9 6.7 Total lactation feed intake, kg 105.15 109.37 5.33 Day 20 of lactation 101.5 102.1 9.3 a,b Means within a row with different superscripts differ (P < 0.01). a,b c,d Means within a row with different superscripts differ (P < 0.01). Means within a row with different superscripts differ (P < 0.05). c,d Means within a row with different superscripts tend to differ *CTL  =  control, and TRT  =  treatment regimen, which provided (P < 0.10). 70% (restriction) and 115% (overallowance) of the CTL diet in the CP and NE contents. The restriction diet was fed from days 28 until 74 *CTL = control, and TRT = treatment regimen, which provided 70 of gestation followed by the overallowance diet for the remainder of (restriction) and 115% (overallowance) of the CTL diet in the CP and gestation. Sows were fed 2.5 kg/d throughout gestation and there were NE contents. The restriction diet was fed from days 28 until 74 of gesta- no refusals. tion followed by the overallowance diet for the remainder of gestation. † † Maximum value. Maximum value. Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/2/2/162/4969850 by Ed 'DeepDyve' Gillespie user on 19 June 2018 166 Farmer et al. end of feeding the RES diet (day 75 of gestation), and protein intake from days 28 until 74 of gestation TRT gilts had less urea, more than double the con- and then overfeeding gilts from days 75 until 110 of centrations of FFA and less IGF-1 than CTL gilts gestation for subsequent lactation performance. An (Table  3, P  <  0.01). After feeding the OVER diet earlier trial suggested that diet deprivation followed (day 110), TRT gilts had more urea than CTL gilts by overallowance during the growing, finishing, (Table  3, P  <  0.01). During lactation, there was a and gestation phases could be beneficial in terms of significant treatment × day interaction for glu - milk yield and mammary gene expression in swine cose (P  <  0.01), a tendency for a treatment × day (Crenshaw et  al., 1989). Yet, a recent experiment interaction for FFA (P < 0.10), and a day effect for showed that when a similar diet deprivation-over- urea (P < 0.01), with values increasing as lactation allowance regimen as described by Crenshaw et al. advanced (Table  3). There were no changes in any (1989) was used in gestation only, there was no measured variables due to treatment on day 3 of increase in mammary development of late-pregnant lactation (P > 0.10), but on day 20 of lactation, gilts (Farmer et al., 2014). It was hypothesized that TRT sows tended to have less FFA and more glu- the level of diet deprivation imposed in early ges- cose than CTL sows (Table 3, P < 0.10). There was tation was too severe for the compensatory growth no change in weekly or total lactation feed intake in late gestation to have any beneficial effects. This of sows across treatments (Table  2; P > 0.10) and was suggested by the 12.9 kg lower BW of treated milk DM, fat, protein, and lactose contents on gilts compared with control gilts on day 110 of ges- day 19 of lactation were not altered by treatment tation (Farmer et al., 2014). In the current trial, the (Table 4; P > 0.10). duration of the diet deprivation was reduced, start- The number of live-born piglets did not vary ing on day 28 of gestation, while an overallowance between treatments, being 13.7 and 12.5  ±  0.5 for similar to previous trials was used in later gestation. CTL and TRT sows, respectively (P  =  0.1). Piglet As expected, concentrations of FFA at the end of BW at birth or on days 1, 7, 14, or 21 of lactation the restricted period were increased due to greater was not affected by treatment, nor was BW gain use of energy reserves (Barb et  al., 1997; Farmer over the lactation period altered (Table 5; P > 0.10). et al., 2014; Ren et al., 2017) and urea was reduced There was no difference between treatments on the indicative of a lower supply of proteins (Pedersen frequency of piglet mortality that occurred either in et al., 2016). The 4.6 kg difference in BW between the first 48 h postpartum (12 vs. 11 sows with dead TRT and CTL gilts on day 110 of gestation in the piglets for CTL and TRT, respectively; P > 0.10) current experiment was much lower than that pre- or over the whole lactation period (7 vs. 10 sows viously seen (Farmer et al., 2014) and was not sig- with dead piglets for CTL and TRT, respectively; nificant. Nevertheless, the increase in BW of gilts P > 0.10), but the incidence of stillborn was greater from days 75 to 110 of gestation was not as large as in CTL than TRT sows (13 vs. 7 sows with stillborn the reduction in BW from days 28 until 75, so that piglets for CTL and TRT, respectively; P < 0.05). even though compensatory growth did take place in DISCUSSION Table 5. Weight of piglets* Groups Findings from the current experiment indicate that there is no beneficial effect of restricting energy Item CTL TRT SEM BW, kg Table  4. Milk composition on day 19 of lactation Birth 1.43 (12.9) 1.50 (12.7) 0.05 (n = 12)* Day 1 of lactation 1.45 (12.3) 1.51 (12.2) 0.05 Day 7 of lactation 2.75 (12.3) 2.80 (11.7) 0.10 Groups Day 14 of lactation 4.52 (11.9) 4.51 (11.7) 0.13 Item CTL TRT SEM Day 21 of lactation 6.10 (11.9) 6.02 (11.7) 0.20 DM, % 18.78 19.52 0.42 BW gain, kg Fat, % 7.30 7.78 0.37 Days 1 to 21 4.65 4.50 0.17 Protein, % 5.38 5.65 0.18 Average numbers of piglets per litter are shown in parenthesis. Lactose, % 4.86 4.71 0.11 *CTL  =  control, and TRT  =  treatment regimen, which provided 70 (restriction) and 115% (overallowance) of the CTL diet in the CP *CTL = control, and TRT = treatment regimen, which provided 70 and NE contents. The restriction diet was fed from days 28 until 74 (restriction) and 115% (overallowance) of the CTL diet in the CP and of gestation followed by the overallowance diet for the remainder of NE contents. The restriction diet was fed from days 28 until 74 of gesta- gestation. tion followed by the overallowance diet for the remainder of gestation. † † Maximum value. Maximum value. Translate basic science to industry innovation Downloaded from https://academic.oup.com/tas/article-abstract/2/2/162/4969850 by Ed 'DeepDyve' Gillespie user on 19 June 2018 Gestation feeding and sow lactation 167 late gestation it was likely not important enough to across treatments are in agreement with the unal- stimulate mammary development. This hypothesis tered milk composition. The similar birthweight of is substantiated by the fact that circulating IGF-1 piglets across treatments also indicates that fetal concentrations were lower in TRT than CTL gilts growth was not affected even though there may on day 75 of gestation and were not greater on day have been a positive effect on the incidence of still- 110 of gestation. Lower IGF-1 concentrations in borns. Yet, the number of animals used was too animals with a slower BW gain were expected due low to determine such an effect and this needs to to the positive relationship between IGF-1 concen- be corroborated in future studies. Ren et al. (2017) trations and growth rate in growing pigs (Owens recently demonstrated that feed restriction or over- et al., 1999). Taking into account the important role feeding of gilts during three short periods of gesta- of IGF-1 for mammary development in rodents tion did not affect litter performance. Even though (Kleinberg and Barcellos-Hoff, 2011) and for mito- growth performance of gilts was increased during genesis of mammary epithelial cells in ruminants those periods when they were fed twice the main- (Forsyth, 1996), one might suggest that for a late tenance requirements, it did not lead to beneficial gestation compensatory feeding strategy to have effects during the subsequent lactation. Therefore, beneficial effects on lactation performance it would altering growth rate of gilts during gestation may need to increase IGF-1 concentrations. not be an adequate strategy to enhance mammary The compensatory feeding regimen was imposed development or the changes incurred need to be in gestation only in the present trial, because when more prominent to have an effect. a feeding regimen similar to that of Crenshaw et al. In conclusion, diet deprivation of gilts as of (1989) was used in the growing-finishing period, day 28 of gestation followed by overfeeding from there was no beneficial effect on mammary devel - day 75 of gestation until farrowing did not improve opment either at puberty (Farmer et al., 2012a) or lactation performance. It is likely that the compen- at 110 d of gestation (Lyvers-Peffer and Rozeboom, satory growth that took place in late gestation was 2001; Farmer et  al., 2012b). In studies by Farmer not adequate to illicit beneficial effects on mam - et al. (2012a, 2012b), the greater BW gain observed mary development because IGF-1 concentrations by Crenshaw (1990) in treated gilts at the end of were not increased. the growing-finishing period was not seen and in the study by Lyvers-Peffer and Rozeboom (2001), LITERATURE CITED BW of gilts was not reported. In fact, gilts sub- Agriculture and Agri-Food Canada. 1993. Recommended code jected to the restricted and overallowance feeding of practice for the care and handling of farm animals - regime weighed 6.2 kg less at puberty than control Pigs. Publ. No. 1898E. Ottawa, ON (Canada): Agriculture and Agri-Food Canada. gilts (Farmer et al., 2012a). Differences in the type AOAC. 2005. Official methods of analysis international. 18th of fiber used and in feed composition were thought ed. Arlington (VA): Assoc. Off. Anal. Chem. to be the cause of this discrepancy. Previous results Barb, C. R., R. R. Kraeling, G. B. Rampacek, and C. R. Dove. (Farmer et  al. 2012a, 2012b) combined with cur- 1997. Metabolic changes during the transition from the rent results therefore question the hypothesis that fed to the acute feed-deprived state in prepuberal and the beneficial effect seen by Crenshaw et al. (1989) mature gilts. J. Anim. Sci. 75:781–789. Crenshaw, J. D. 1990. Feeding gilts to enhance lactational per- on sow milk yield was mostly due to compensa- formance. Fargo (ND): North Dakota State Univ. Swine tory growth in late gestation. It may be that a feed- Res. Rep. Agric. Exp. Stn. Ext. Services. p.12–23. ing regime that could bring about the 8.2  kg BW Crenshaw, J. D., C. S. Park, P. M. Swantek, W. L. Keller, and R. advantage of treated gilts at the end of the grow- C. Zimprich. 1989. Lactation response of gilts to a phased ing-finishing period in Crenshaw’s study would be feeding regimen designed to induce compensatory growth. J. Anim. Sci. 67(Suppl. 2):107–108. (Abstr.) beneficial for mammary development and future Farmer, C., M. F. Palin, and Y. Martel-Kennes. 2012a. Impact milk yield. However, circulating concentrations of of diet deprivation and subsequent over-allowance during IGF-1 were not measured in the study by Crenshaw prepuberty. Part 1. Effects on growth performance, metab- et al. (1989), so it is unknown if the beneficial effect olite status, and mammary gland development in gilts. J. on sow milk yield was mediated through this growth Anim. Sci. 90:863–871. doi:10.2527/jas.2011-4131 factor. Farmer, C., M. F. Palin, and Y. Martel-Kennes. 2012b. Impact of diet deprivation and subsequent over-allowance during The lack of treatment effect on circulating prepuberty. Part 2.  Effects on mammary gland develop- concentrations of IGF-1 on day 110 of gestation, ment and lactation performance of sows. J. Anim. Sci. the presence of only tendencies for decreased FFA 90:872–880. doi:10.2527/jas.2011-4480 and increased glucose in TRT vs. CTL gilts on that Farmer, C., M. F. Palin, and Y. Martel-Kennes. 2014. 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Journal

Translational Animal ScienceOxford University Press

Published: Apr 13, 2018

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