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Dietary B vitamin needs of strains of pigs with high and moderate lean growth^1,2

Department of Animal Science, Iowa State University, Ames 50011

	Abstract

Top Abstract INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION LITERATURE CITED

 
Ten sets of 5 littermate pigs from each of 2 genetic strains were utilized to determine the impact of the dietary concentration of 5 B vitamins (riboflavin, niacin, pantothenic acid, cobalamin, and folacin) on growth from 9 to 28 kg of BW in pigs with high or moderate capacity for lean growth. All pigs (penned individually) were reared via a segregated, early weaning scheme, so that the lean growth potential of each strain could be expressed. The basal diet provided the 5 test vitamins at concentrations of total and estimated bioavailability equivalent to a minimum of 100 and 70%, respectively, of their estimated requirements (NRC, 1998) for 5- to 10-kg pigs. At a BW of 9 ± 0.9 kg, pigs within each litter were allotted to the basal diet supplemented with sources of the 5 test vitamins equivalent to an additional 0, 100, 200, 300, or 400% (bioavailable) of the NRC requirements. Pigs from the high lean strain consumed less feed (P < 0.05) and gained BW faster (P < 0.02) and more efficiently (P < 0.01) than pigs of the moderate lean strain. In both lean strains, the rate and efficiency of growth were improved (P < 0.01) as dietary B vitamin concentrations were increased. However, the dietary B vitamin concentrations needed to optimize G:F were greater (P < 0.03) in the high (>470% of NRC, 1998) vs. moderate (270%) lean strain. Based on these data, the dietary needs for 1 or more of the 5 B vitamins are greater than current NRC (1998) estimates, particularly in pigs expressing a high rate of lean tissue growth. The greater need for these vitamins is not associated with greater dietary energy intake or body energy accretion rate but is potentially due to shifts in the predominant metabolic pathways.

Key Words: B vitamin • lean growth strain • pig

	INTRODUCTION

Top Abstract INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION LITERATURE CITED

 
Current estimates of dietary vitamin requirements for swine (NRC, 1998) are largely based on studies conducted 20 to 50 yr ago. For young pigs, the estimated B vitamin requirements are based in part on the following work for niacin (Braude et al., 1946; Firth and Johnson, 1953; Harmon et al., 1963), pantothenic acid (Strothers et al., 1955; Barnhart et al., 1957; Sewell et al., 1962; Harmon et al., 1963; Palm et al., 1968), riboflavin (Krider et al., 1949; Miller and Ellis, 1951; Harmon et al., 1963), vitamin B₁₂ (Anderson and Hogan, 1950; Catron et al., 1952), and folacin (Easter et al., 1983).

Because of changes in industry practices during the last 20 to 50 yr, vitamin requirements for swine now are potentially greater. Modern genetic strains of pigs have a greater capacity for body growth, particularly proteinaceous (lean) tissue accretion. Furthermore, use of management techniques such as all-in-all-out production and segregated weaning schemes minimize the levels of immune system activation the pig experiences, which further increases the animal’s capacity for body protein accretion (Williams et al., 1997a). These low immune status pigs from high lean strains can deposit 12 to 18 g of protein per unit of metabolic BW (BW^0.75) daily, compared with 4 to 9 g for fat and lean genetic strains experiencing moderate to highly stimulated immune systems that were evaluated in the 1950s through the 1970s (Stahly, 1986; Williams et al., 1997a; Frederick and Stahly, 1999). Assuming the vitamin needs per unit of body protein accretion are relatively constant, we hypothesized that pigs with high capacities for lean tissue growth would require 2 to 4 times the daily B vitamin needs currently defined by NRC (1998).

The objective of this study was to determine the impact of the dietary concentration of a group of 5 B vitamins on the rate and efficiency of growth of pigs with a low level of immune system activation and with a high or moderate genetic capacity for lean tissue accretion.

	MATERIALS AND METHODS

Top Abstract INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION LITERATURE CITED

 
All procedures involving animals were approved by the Institutional Animal Care Committee.

Experimental Treatments
The experimental treatments consisted of 2 lean growth genetic strains and 5 dietary concentrations of a group of 5 B vitamins. The 2 lean growth strains consisted of sources of pigs with a high or a moderate genetic capacity for lean tissue accretion. The daily lean tissue growth capacities of the high and moderate lean growth genotypes are estimated to be 0.35 to 0.39 and 0.25 to 0.30 kg, respectively, from 18 to 110 kg of BW. These capacities are based on previous measures of body growth rates and carcass traits of the 2 genetic sources at our research station (data not shown).

The dietary regimens consisted of a basal diet supplemented with 5 levels of specific B vitamins. The basal diet consisted of (as-fed basis) a mixture of corn-soybean meal, 15% lactose, 5% casein, and 5% choice white grease (Table 1)calculated to contain 3.422 Mcal of ME/ kg and 1.8% lysine. The diet was formulated to meet or exceed the pig’s requirements for all nutrients except for the 5 test vitamins. The 5 test vitamins, niacin, pantothenic acid, riboflavin, B₁₂, and folacin, were chosen because they are closely involved in fueling proteinaceous tissue accretion via support of energy metabolism and synthesis of methyl groups and nucleic acids.

The synthetic sources used for the 5 test vitamins were nicotinic acid (99.5%), riboflavin (96%), D-calcium pantothenate (92% pantothenic acid), cyanocobalamin (1%), and folic acid (100%), respectively. The bioavailability of vitamins in each of the supplemental vitamin sources was assumed to be 100%. In the experimental diets, synthetic sources of each of the 5 test vitamins were added to the basal diet at the expense of cornstarch to provide dietary concentrations of bioavailable vitamins equivalent to 70, 170, 270, 370, and 470% of NRC (1998) estimated requirements for 5- to 10-kg pigs for this group of B vitamins. All other vitamins were provided at levels that met or exceeded 600% of the estimated requirements (NRC, 1998) for 5- to 10-kg pigs to ensure they were not limiting growth. A single source of each ingredient was used to eliminate variation in vitamin content.

The analyzed concentration of the 5 test vitamins in the experimental diets is reported in Table 2.These values represent analyzed total and bioavailable (analyzed total content corrected for the estimated bioavailability of vitamins in each ingredient) B vitamin contents. Niacin, pantothenic acid, riboflavin, and B₁₂ concentrations were analyzed by AOAC (1990) procedures, and folacin was analyzed by MAIF (1985) procedures (Hazleton Laboratories, Madison, WI).

From 12 to 16 d of age, pigs were self-fed a pelleted, milk-based diet containing no supplemental vitamins. From d 16 postweaning to 9 kg of BW, pigs were selffed the basal diet. At a BW of 9 ± 0.9 kg, pigs were randomly allotted within each litter to 1 of the 5 dietary vitamin regimens, resulting in 10 pens/dietary B vitamin concentration for each strain. Pig weights and feed consumption were measured at 4-d intervals until each animal reached a BW of 29.5 ± 2.3 kg. Three pigs were removed from test (1 high lean strain on 270% B vitamins; 2 moderate lean strain—1 on 70% and 1 on 270% B vitamins).

Immune status of the pigs was monitored at BW of 9 (initial), 19, and 28 (final) kg via microfluorometric determination of the proportion of lymphocytes expressing CD₄ and CD₈ surface cell antigens (Williams et al., 1997b,c). Serological titers for prevalent antigens in the pigs were analyzed by the Iowa State University Diagnostic Laboratory for Actinobacillus pleuronpneumoniae, mycoplasma hyopneumonia, porcine reproductive and respiratory syndrome, and swine influenza virus, as outlined by Williams et al. (1997b, c).

Data were analyzed as a split-plot design (Steel and Torrie, 1980) using the GLM procedure (SAS Inst. Inc., Cary, NC). Lean growth strains were considered the whole-plot treatments, and dietary B vitamin concentrations were considered the subplot treatments. Responses of pigs to dietary vitamin concentrations at different stages of the pig’s development were analyzed as repeated measures. The pig was considered the experimental unit. Least square means are reported. The error terms of replicate(strain) and replicate x B vitamins(strain) were used to test strain and B vitamin effects, respectively. Linear, quadratic, and cubic contrasts were used to determine the responses to increasing concentrations of dietary B vitamins.

	RESULTS AND DISCUSSION

Top Abstract INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION LITERATURE CITED

 
Diet Composition
The analyzed vitamin content of the experimental diets approximated the initial calculated values for niacin, pantothenic acid, riboflavin, and B₁₂ but were higher for folacin (Table 2). The analyzed vitamin content in the basal diet expressed as a percentage of the initial calculated values were 107, 93, 118, 107, and 203%, respectively, for niacin, pantothenic acid, riboflavin, vitamin B₁₂, and folacin. The incremental additions of the 5 test vitamins analyzed to be 103, 97, 105, 94, and 147%, respectively, of the projected additions [calculated as increments of 100% NRC (1998) for 5- to 10-kg pigs].

Immune Status of Experimental Animals
The experimental protocol required that the experimental animals experience a low antigen challenge in order for the pigs to express a high rate of proteinaceous tissue growth and thus nutrient needs. Furthermore, minimizing the animals’ antigen exposure was required in order for the genetic capacities of the 2 genotypes to be expressed largely independent of confounding environmental factors (i.e., pig health). Based on analysis of serological titers for common antigens as well as lymphocyte surface markers, the experimental animals from both genetic strains experienced a low level of antigen exposure and thus a low level of immune system activation throughout the study (Table 3).The low proportion of peripheral blood lymphocyte expressing CD4+ surface markers also indicates that the pigs in both strains experienced a low level of antigen exposure and a low level of immune activation. In the current study, the CD4+/CD8+ ratios ranged from 0.62 to 0.83 which are in close agreement with values of 0.6 to 0.8 previously observed in low antigen exposed, segregated early weaned-reared pigs at similar stages of growth (Williams et al., 1997b,c). In pigs experiencing chronic antigen exposure, the CD4+/CD8+ ratios are greater, averaging 1.8 to 2.2. Lower expressions of CD4+ surface markers and CD4+/CD8+ ratios are associated with greater feed intakes and improved BW gains, gain/feed ratios, and DM and protein digestibilities in pigs (Williams et al., 1997a,b,c).

Strain x B Vitamins x Pig Weight Effects
The daily BW gains and gain:feed ratio of pigs during 4-d periods when each pig reached BW (±2.25 kg) of 9.75, 14.25, 18.75, 23.25, and 27.75 were analyzed. Daily feed intake and daily BW gains increased (P < 0.01) as pigs grew from 9 to 28 kg of BW. Efficiency of feed utilization was reduced (P < 0.01) as the pigs matured. Based on efficiency of feed utilization, the high lean growth pigs weighing 9.75 to 14.25 kg and 18.75 to 27.75 kg, respectively, needed dietary B vitamin concentrations equivalent to 470% or greater and 370% of the NRC (1998) requirements for 5- to 10-kg pigs (Table 5). In moderate lean growth strains, pigs weighing 9.75 to 14.25 kg and 18.75 to 27.75 kg needed dietary B vitamin concentrations equivalent to 370 and 270% of the NRC (1998) requirements for 5- to 10-kg pigs (Table 5). Because the estimated B vitamin requirements, expressed as milligrams per kilogram of diet, decline as animals grow (NRC, 1998), the high lean growth pigs at 14 and 28 kg of BW were actually responding to B vitamin concentrations equivalent to about 600 and 500% of their estimated needs for their respective BW.

At the dietary vitamin concentrations needed to optimize efficiency of feed utilization at each stage of growth, the daily intakes of estimated bioavailable vitamins were greater in the high lean strain even though their daily intakes of feed, and thus energy, were less. For example, daily intakes of estimated bioavailable riboflavin associated with optimum gain/feed ratios were 0.6, 1.9, 3.9, 4.8, and 6.1 mg/pig greater in the high vs. moderate lean strain at mean BW of 9.75, 14.25, 18.75, 23.25, and 27.75 kg, respectively. These greater daily vitamin intakes also were associated with greater daily BW gains in the high vs. moderate lean strains at each stage of growth. At vitamin concentrations needed to optimize gain:feed, the amount of bioavailable vitamin that supported a kilogram of BW gain was relatively constant among pig strains and stages of growth. For example, the milligrams of bioavailable riboflavin consumed per kilogram of BW gain averaged 23 (range of 20 to 24 for the 5 stages of growth) and 21 (range of 19 to 24) for the high and moderate strains, respectively. In contrast, only 6 mg of riboflavin/kg of BW gain is estimated to be needed by the NRC (1998) based on work in pigs that were accruing BW less rapidly and in which a lower proportion of the accrued tissue was lean.

In Table 6, the performance of the lean, high health pigs in the current study are compared with that estimated by NRC (1998) for similar BW pigs. The high and moderate lean strains in the current study exhibited 19 and 8% greater daily BW gains and required 31 and 18% less feed to achieve each unit of BW gain than that of pigs for which NRC (1998) vitamin requirements are defined. Based on body composition of high and moderate lean strains of segregated early weaned-reared pigs from previous studies at our station (Lutz and Stahly, 1998; Stahly and Lutz, 2000), the composition of BW gain also differs among the 3 pig groups. The high and moderate lean strains in the current study are estimated to accrued daily 46 and 30% more body protein, 50 and 51% less body fat, and 27 and 32% less body energy, than that estimated by NRC (1998). Although the pigs in the current study were accruing less body energy daily, these animals were partitioning twice the proportion of their consumed energy (ME) into body protein accretion than that estimated by NRC (1998). Specifically, the high lean strains are estimated to use 41 and 32%, respectively, of their consumed energy for body protein and fat accretion compared with 21 and 53%, respectively, as estimated by the NRC (1998).

The need for greater amounts of some B vitamins in high lean, high health animals in the current study could be due to shifts in the metabolic profile of these animals. These shifts include 1) greater amounts of muscle growth, particularly the rapidly proliferating and differentiating satellite muscle cells (Mesires and Doumit, 2002; McCroskery et al., 2003; Steelman et al., 2006); 2) greater predominance of glycolytic metabolism due to an increased predominance of glycolytic vs. oxidative muscle fibers (Lefaucheur et al., 2004) and greater number of satellite (low mitochondria, predominant glycolytic metabolizing) muscle cells (Barani et al., 2003); and 3) greater proportional metabolic demand for muscle accretion vs. body maintenance, immune functions, or fat tissue accretion. Additional differences could potentially include greater insulin sensitivity due to lower body fat accretion, lower rates of fat mobilization, and lower proportions of dietary fat bypassing oxidative processes and being directly deposited in body fat stores. In lean strains, muscle accretion rates are increased, which are strongly associated with greater rates of satellite cell proliferation and differentiation (Mesires and Doumit, 2002). Furthermore, the predominance of glycolytic muscle fibers in pigs has been shown to increase significantly as the animal’s capacity to accrue body protein is increased due to genetic selection for lean tissue growth (Lefaucheur et al., 2004), reduction of antigen exposure (Tiao et al., 1997), minimization of physical exercise (McAllister et al., 1997; Gondret et al., 2005), optimization of the thermal climate (Lefaucheur et al., 1991), and use of growth-enhancing compounds (Depreux et al., 2002). This greater predominance of glycolytic metabolism in cells is normally associated with a diminished capacity for ATP production, minimization of the Krebs cycle in energy processing, and a greater dependence on glucose as an energy source as well as a provider of needed precursors (i.e., ribose and NADPH) for nucleotide and lipid biosynthesis (Murray et al., 2003). In recent work with other cells (i.e., cancer) expressing a high degree of glycolysis and a rapid rate of proliferation, similar responses of diminished capacities for ATP production, reduced Kreb cycle activities, and a greater dependence on glucose are observed (Perumal et al., 2005a,b). Oral administration of additional riboflavin and niacin in rats experiencing mammary carcinoma has been shown to largely restore these cells’ energy production and Krebs cycle activities to near normal levels, which indicates a higher metabolic need for energy modulating vitamins in these cells (Perumal et al., 2005a,b). The impact of this greater predominance of glycolytic muscle cells/tissues in modern lean strains of high health pigs on the response of these animals to specific energy modulating vitamins has not been quantified.

Because maintenance processes would represent a smaller proportion of the metabolic demands in high health, rapidly growing animals, their proportional needs for methyl groups (i.e., methionine synthesis, intestinal cell turnover, immune cell proliferation, regeneration of endogenous antioxidant capacity), as well as denovo synthesized nucleic acids to support body maintenance and immune system functions, would be expected to be less. Thus, the metabolic demand for vitamin B₁₂, folacin and possibly niacin for these functions in high health animals would be less; however, their total need could be increased in rapidly growing animals due to a potentially large amount of nucleotide synthesis in proliferating satellite muscle cells.

Based on these data, the greater capacity for lean strains of high health pigs to efficiently accrue BW, particularly proteinaceous tissue, results in a greater dietary need for 1 or more of the test B vitamins. The greater need for these vitamins in the lean, high health pig is not associated with greater dietary energy intake or body energy accretion rate but potentially is due in part to shifts in the predominant metabolic pathways employed.

	Footnotes

 
¹ Journal paper No. J-19153 of the Iowa Agriculture and Home Economics Experiment Station, Ames, Iowa, Project No. 3142, and supported by Hatch Act and State of Iowa funds.

² Research supported in part by BASF Corporation, Mount Olive, NJ.

³ Department of Animal Science, 201 Kildee Hall, Iowa State University, Ames, IA 50011.

⁵ Current address: Sygen Inc., Franklin, KY.

⁶ Current address: McFleeg Feeds, PO Box 1205, Watertown, SD 57201.

⁴ Corresponding author: tstahly{at}iastate.edu

Received for publication February 15, 2006. Accepted for publication July 19, 2006.

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This Article Abstract Full Text (PDF) Alert me when this article is cited Alert me if a correction is posted Services Similar articles in this journal Similar articles in PubMed Alert me to new issues of the journal Download to citation manager Citing Articles Citing Articles via Google Scholar Google Scholar Articles by Stahly, T. S. Articles by Swenson, S. G. Search for Related Content PubMed PubMed Citation Articles by Stahly, T. S. Articles by Swenson, S. G.