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Anim Biosci > Volume 35(3); 2022 > Article
Cho, Macelline, Wickramasuriya, Shin, Kim, Son, and Heo: Moderate dietary boron supplementation improved growth performance, crude protein digestibility and diarrhea index in weaner pigs regardless of the sanitary condition



The study was conducted to investigate the impact of boron supplementation on nutrient digestibility, inflammatory responses, blood metabolites and diarrhea index, and their relevance to growth performance in weaned pigs housed in good and poor sanitary environments for 14 days after weaning.


A total of 108 male pigs (Duroc×[Yorkshire×Landrace]) weaned at 21 days of age were used in a randomized complete block design with 2×3 factorial arrangement. Pigs were assigned to three boron treatments (0, 5, and 10 mg/kg) under two environments (good and poor sanitary) to give six replicates per treatment (3 pigs per replicate). On 0, 7, and 14 days, one pig per replicate was euthanized to collect, ileum tissue samples, and rectal fecal samples.


Boron supplementation quadratically influenced (p<0.001) feed intake and weight gain in pigs housed in good sanitary conditions from 1 to 14 days post-weaning where pigs offered 5 mg/kg boron optimized weight gain and feed intake. There is a quadratic interaction (p = 0.019) on feed intake for 1 to 14 days post-weaning where 5 mg/kg boron increased feed intake in good sanitary conditions. Pigs housed in the poor sanitary environment decreased (p<0.001) villus height and crypt depth in ileum at days 7 and 14. On day 7 and 14, crude protein digestibility was quadratically influenced (p<0.05) by boron supplementation. Boron supplementation linearly increased (p<0.05) plasma calcium and cholesterol levels whilst linearly (p = 0.005) reducing plasma triglyceride concentrations. Diarrhea index was quadratically influenced (p<0.05) by boron supplementations regardless of sanitary conditions where 5 mg/kg boron inclusion achieved the lowest diarrhea index.


Pigs offered 5 mg/kg of boron increased weight gain which may be deduced by improved dry matter, crude protein, and energy digestibility regardless of the sanitary conditions.


Boron has been identified as an important element in animals because of its impact on metabolic responses, energy utilization and homeostatic mechanisms in animal tissues [1]. It has been found that boron supplementation improved the growth performance in pigs may be due to its metabolic functions and positive impact on nutrient absorption in the small intestine as described by Armstrong et al [2,3], respectively. Moreover, the beneficial impact of dietary boron on bone strength was discussed in Chen et al [4] such that boron is an indispensable mineral in bone development, cell proliferation and mineralization in bone tissues. Moreover, it has been stated that boron-supplemented diets have increased bone yield stress and bending moment in pigs [5]. Armstrong et al [2] demonstrated that a diet supplemented with 5 ppm boron improved feed conversation efficiency in pigs. Subsequent experiments [46] revealed that pigs fed boron-supplemented diets showed higher growth rates, increased plasma mineral concentrations, and serum metabolite concentrations than pigs fed diets without boron. However, high boron contents in diets adversely impact growth performance and immune organs (e.g., thymus, spleen, and bursa) in animals which is dependable on the type of animal and their body weights (BWs) [7].
Pigs housed in facilities with poor sanitary conditions exhibit an increased incidence of inflammatory disease that leads to poor growth performance [8]. Proinflammatory cytokines play a major role in the animal immune system under inflammatory disease conditions by stimulating the possible defensive mechanism and inflammatory responses [9]. Supplementation of antimicrobial growth promoters (AGP) has long been used to overcome enteric challenges but presently there are many limitations for AGP usage for the pig industry mainly due to antimicrobial resistance and environmental pollution [10]. Interestingly it has been found that boron has the ability to improve nutrient digestibility by moderating nutrient transporters and stimulate the immune system by producing pro-inflammatory cytokines [16]. Therefore, boron may be a suitable candidate for replacing AGP in pigs. Testing the impact of boron for pigs under poor sanitary conditions may help to understand its beneficial influences in challenging conditions. In previous studies [2,3,5,6], pigs were exposed to 5 to 15 mg/kg of dietary boron. However, as a safe margin, 5 and 10 mg/kg levels of boron were tested in the present study since a higher concentration of dietary boron may cause toxicity in animals [7]. Consequently, this study was conducted to test the hypothesis that dietary boron supplementation improves growth performance and nutrient digestibility with improved inflammation responses and diarrhea index values in pigs housed in an environment with poor sanitary conditions for 14 days post-weaning.


The experimental protocol used in this study was approved by the Animal Ethics Committee of the Chungnam National University (CNU-00485).

Experimental design, animals, housing and diets

This experiment was conducted as a 2×3 factorial treatment arrangement, with respective factors being i) two environmental conditions (good sanitary, cleaned and disinfected previously unpopulated rooms; poor sanitary, manure application with no cleaning or disinfection of a previously populated room) and ii) three levels of boron (0, 5, and 10 mg/kg).
A total of 108 male pigs (Duroc×[Yorkshire×Landrace]) weaned at 21 days of age with initial BW of 6.59±1.86 kg were used. Pigs were obtained from a local government experimental farm (Chungnam Livestock Research Institute, Cheongyang, Korea) at weaning and transported to an animal research farm. The facility contained two rooms that allowed pigs to be housed separately in good sanitary or poor sanitary environments, to avoid any cross contamination. Pigs were allocated to their experimental treatments based on initial BW for blocking the factors within two rooms (randomized complete block design). Six replicate pens with three pigs per pen were allocated to each treatment. Each pen was equipped with a nipple bowl drinker and a metal trough. The ambient temperature was maintained at 29°C± 1°C for the initial week and then gradually decreased by 1°C every week. Pigs were offered the experimental diets on an ad libitum basis for 2 weeks and were always freely accessible to the freshwater.
Experimental diets were formulated to have similar crude protein and gross energy levels and meet or exceed the NRC [11] recommendations (Table 1). Three levels of boron (0, 5, and 10 mg/kg) were included by supplementing boric acid (Junsei Chemical Co., Ltd., Tokyo, Japan) by top dressing. Chromic oxide was added to experimental diets (0.3%) as an indigestible marker for the calculation of apparent total tract digestibility (ATTD) of dry matter (DM), crude protein, and energy.

Measurement of BW and feed consumption

Individual BW was measured at days 0, 7, and 14 of the experiment. Feed intake of each pen was recorded on days 7 and 14 as feed disappearance from the feeder. Based on the measurements, the average daily gain (ADG, g/pig/d), average daily feed intake (ADFI, g/pig/d) and gain-to-feed (G:F) ratio were calculated.

Assessment of fecal consistency and the incidence of diarrhea

Two weeks after weaning, feces were visually assessed daily at 1000 h to determine fecal consistency scores and the incidence of diarrhea using the procedure described by Heo et al [12] during the experimental period (0 to 14 days). Feces were assessed using the fecal consistency score according to Marquardt et al [13] using a subjective score on a 3-point scale ranging from 1 to 3: 1, well formed; 2, sloppy; 3, diarrhea. On days 7 and 14, fecal samples were collected from the rectum and stored in a labeled sterile container and kept frozen at −20°C until analyses for DM, crude protein, and gross energy.

Post-mortem procedure

One pig per pen was euthanized on days 0, 7, and 14 according to modified procedures described in [13] to collect blood and intestinal tissue samples. Pigs were administered a single intramuscular injection of 0.1 mL SUCCIPHARM (50 mg suxamethonium chloride, Komipharm International Co., Ltd., Shieung, Korea) to induce general anesthesia, and then blood samples (5 to 10 mL) were collected via jugular vein puncture using vacuum tubes coated with lithium heparin (Vacutainer; Becton Dickinson, Franklin Lakes, NJ, USA). The blood plasma was separated by centrifugation (Micro 12; Hanil Science Co., Ltd., Incheon, Korea) at 2,000×g for 10 min at 4°C and stored at −80°C until analysis. Thereafter, pigs were introduced into a chamber where residual air was rapidly flushed with 100% CO2 until death was confirmed. The abdominal cavity was exposed by midline laparotomy. For measurement of villous height and crypt depth, 3 to 4 cm section segment of the small intestine was removed at the ileum (approximately 5 cm cranial to the ileo-caecal junction) as described by Heo et al [14] and carefully washed with phosphate buffered saline and preserved in 10% formalin solution for subsequent histological examination.

Mucosal histology

Tissue preparation and methods for microscopic measurements were conducted following standard histological procedures described by Heo et al [14]. After fixation in the 10% phosphate buffered saline for several days, ileum sections were excised, dehydrated, and embedded in paraffin wax. From each of the embedded samples, 6 transverse sections (4 to 6 μm) were cut, stained with hematoxylin and eosin, and mounted on glass slides. The height of 10 well oriented villi and their associated crypts were measured with a light microscope (OLYMPUS CX31, Tokyo, Japan) using a calibrated eyepiece graticule [15].

Chemical analyses

Diet DM concentration was determined according to the AOAC 930.15 [16] by oven drying 5 g of samples at 135°C for 2 h. The gross energy content of experimental diets and feces were measured using an isoperibol bomb calorimeter (model 6300; Parr Instrument, Moline, IL, USA) which has been calibrated using benzoic acid as a standard. Nitrogen contents in the experimental diets and feces were determined by the combustion method of AOAC 990.03 [16] using the LECO nitrogen analyzer (model CNS-2000; LECO Corp., St. Joseph, MI, USA) and crude protein was calculated as N×6.25.
Plasma samples were used to quantify the concentrations of proinflammatory cytokines by using commercially available ELISA kits (R&D Systems, Minneapolis, MN, USA) for interleukin 1β (IL-1β) and tumor necrosis factor-α (TNF-α) according to the manufacturer’s instructions following the method described by Piñeiro et al [17].

Digestibility calculation

The ATTD of DM, crude protein, and gross energy were calculated for each diet according to Stein et al [18] by using the following equation.
ATTD (%)=1-(Nutrient in feces/Nutrient diet)×(Cr in diet/Cr in feces)×100%

Statistical analyses

The pen was the experimental unit for all responses. Data were analyzed as randomized complete block design using the general linear model procedure of SPSS software (Version 22; IBM SPSS 2013). Orthogonal polynomial contrast was performed to get the linear and quadratic effect of dietary boron and interaction effects on response parameters. Pigs were blocked based on weaning weight and the block was used as a random factor in the model for all measured experimental variables. Body weight, ileum morphology and immune response on day 0 were included in the model as a covariate for analyses of growth performance, ileal architecture and cytokine responses. Data for the incidence of post-weaning diarrhea were expressed as the mean percentage of days with diarrhea relative to the 14 days after weaning [12].


Growth performance

The effects of dietary boron and environmental condition on growth performance in weaned pigs were presented in Table 2. There was no mortality found through the experiment period. There was a quadratic interaction on ADFI for 1 to 7 (p = 0.009) and 1 to 14 (p = 0.019) days where pigs supplemented 5 mg/kg attained the highest feed intake under good sanitary conditions but not in the poor sanitary conditions. During 8 to 14 days, a linear interaction effect was found on G:F ratio where the transition of dietary boron from 0 to 10 mg/kg linearly reduced G:F ratio (p = 0.049). Boron supplementation quadratically influenced ADG (p< 0.001) and G:F (p = 0.014) in pigs from 1 to 14 days post-weaning regardless of sanitary conditions.

Intestinal morphology

The impact of environmental conditions and dietary boron levels on ileal morphology in weaned pigs was tabulated in Table 3. There was no interaction effect found on ileal morphology during the experimental period. Pigs housed in good sanitary conditions obtained higher villus height, crypt depth and lower villous height to crypt depth ratio (V:C) than their counterpart (p<0.01).

Apparent digestibility of nutrients

Effects of environmental condition and dietary boron levels on ATTD of DM, crude protein, and energy in weaned pigs shown in Table 4. On day 7, the interaction effect indicates that crude protein digestibility linearly reduced by supplemented boron in good sanitary conditions (p = 0.013). Good sanitary conditions improved DM, crude protein, and energy digestibility compared to poor sanitary conditions (p<0.05). Boron supplementation quadratically influenced DM, crude protein, and energy digestibility at day 7 and 14 (p<0.05).

Pro-inflammatory cytokines

Table 5 showed the effect of environmental conditions and boron levels on pro-inflammatory cytokines in the gut mucosa of weaned pigs on day 7 and 14. Neither interaction nor main treatment effect was found on pro-inflammatory cytokine levels in the present study.

Plasma composition

Plasma concentrations of calcium, phosphorous, cholesterol and triglyceride at day 7- and 14-days post-weaning were presented in Table 6. There were linear (p = 0.014) and quadratic (p = 0.031) interactions obtained on plasma cholesterol concentrations on day 7. Sanitary conditions influenced plasma phosphorus levels on day 7 and 14 (p<0.001). On day 7, supplemented boron linearly increased plasma calcium levels (p = 0.026) but linearly reduced phosphorous (p<0.001) and triglyceride (p<0.001) levels. On day 14, supplemented boron linearly increased plasma calcium (p = 0.004) and cholesterol levels (p = 0.037) whereas reduced triglyceride levels (p<0.005).

Diarrhea index

Impacts of environmental conditions and supplemented boron levels on diarrhea index in weaned pigs are tabulated in Table 7. No interaction effect was observed (p>0.05); otherwise, prominent environment effect obtained for up to 7 days post-weaning where pigs housed in the poor sanitary conditions obtained higher (p = 0.05) diarrhea index than their counterpart. Supplemented boron quadratically influenced diarrhea index for 1 to 7 days (p = 0.022), 8 to 14 days (p = 0.020), and 1 to 14 days (p = 0.002) where lower diarrhea index was supported by the pigs offered 5 mg/kg supplemented diets.


It has been found that pigs fed diets with 5 mg/kg boron supplementation improved growth performance in pigs under conventional farm conditions [2]. In the present study, the quadratic effects of boron supplementations on growth performance indicative that 5 mg/kg boron supplementation supported higher feed intake, weight gain, and G:F ratio in weaner pigs. However, the transition of boron supplementation from 0 to 10 mg/kg reduced feed intake by 35% (442 vs 289 g/pig/d, p = 0.006) and 35% (380 vs 245 g/pig/d, p = 0.022) in poor sanitary and good sanitary conditions, respectively, where the significance of pair-wise comparisons are shown in parentheses for 1 to 14 days post-wean. It has been reported that dietary boron can be toxic to animals by damaging immune organs; consequently, decreased feed intake and reduced weight gain [7]. Therefore, 10 mg/kg of boron supplementation may have become toxic regardless of sanitary conditions for weaner pigs in the present study. Consequently, boron supplementation at 5 mg/kg may be the safe margin in weaner pigs which is needed to optimize their growth performance. However, the quadratic equation obtained for weight gain (YADG = 363+42.8X(boron)− 5.76X2(boron); r = 0.632, p<0.001) showed that 3.71 mg/kg level of boron supplementation attained the highest ADG of 442 g/pig/d for 1 to 14 days post-wean in the present study.
It has been reported that boric acid supplementation with water (640 mg/L) stimulates intestinal cell apoptosis in poultry and resulted from higher intestinal cell proliferation as a compensative reaction [7]. Moreover, intestinal cell apoptosis contributed to cell proliferation could results lower V:C ratio due to higher demand for intestinal cell turnover to maintain the villus physiology. However, it was unable to detect such an impact in intestinal morphology from dietary boron supplementation in the present study. It has been reported that Pigs reared under unclean sanitary conditions had shorter villus height and less crypt depth in Zhao et al [19] which is similar to the present study. Reduced villus height and crypt depth were resulted due to the higher pathogenic bacteria activity and their products in the gastrointestinal tract as reported by Ao et al [20]. Pigs in poor sanitary conditions have a higher possibility to contain a large number of pathogenic bacteria in their gastrointestinal tract and it could be the reason for impaired gut morphology obtained in the present study. However, impaired intestinal morphology was not reflected in growth performance in the present study. The possible explanation for these outcomes may be due to the reporting morphology of ileum instead of jejunum, thus jejunum is the major site where the majority of nutrient absorption takes place whilst most of the bacterial activity takes place in the ileum.
The quadratic effect of dietary boron supplementation indicated that 5 mg/kg boron improved crude protein digestibility on day 7 and 14. Goldbach et al [21] revealed that boron can bind with protease enzyme, which may reduce protein digestibility in animals. Consequently, extensive protease inactivation may be the reason for low crude protein digestibility in pigs offered 10 mg/kg boron diets in the present study. Therefore, low weight gain and G:F ratio in pigs offered 10 mg/kg may be also influenced by poor crude protein digestibility. Suiryanrayna and Ramana [22] reported that lowering gastric pH using weak acids increased protein utilization in pigs by converting inactive pepsinogen to pepsin. In the present study, boron was supplemented as boric acid which is known as weak acid with a pKa value of 9.2 [23] and it also helps to reduce the acid binding capacity in feed. Therefore, the negative impact of boron on protease enzymes may be overcome by the acidity of boric acid when supplementary boron concentration is low (5 mg/kg).
Boron impact on the production of pro-inflammatory cytokinin in pigs is variable in previous studies [3,6]. However, Armstrong and Spears [6] stated that pigs offered boron supplemented diets increased the production of TNF-α under lipopolysaccharide challenge conditions and the mechanism is retained unclear. Pro-inflammatory cytokinin production is important for stimulating the immune system to take place necessary immune responses to protect the body from antigens. Lipopolysaccharides are a type of endotoxins were existing in the membranes of gram-negative bacteria. Poor sanitary conditions influenced pathogenic bacteria activity in the gastrointestinal tract in pigs may reasoning endotoxin leakage to systemic plasma. Therefore, in the present study, it was predicted that dietary boron may influence pro-inflammatory cytokinin production in pigs housed in poor sanitary conditions. However, the outcomes of the present study did not support this hypothesis because both dietary boron and sanitary conditions did not influence the pro-inflammatory cytokinin levels in systemic plasma. Furthermore, it is important to measure plasma endotoxin levels to confirm the endotoxin leakage from the gastrointestinal tract in future studies.
It has been found that boron can influence the utilization and metabolism of calcium, phosphorous, cholesterol, and triglycerides in both humans and animals [7]. In the present study, dietary boron supplementation linearly elevated plasma calcium and phosphorus levels at day 7 and a similar pattern was observed on day 14. Increasing plasma calcium levels may be indicative of improved calcium absorption in the gastrointestinal tract. The evidence for improved calcium absorption with boron supplementation has been reported in Bhasker et al [24] and boron influence on modification of calcium transporters across plasma membranes was reported in Green [25]. As a confounding factor, the acidity of the boric acid used in the present study may be also influenced the calcium and phosphorus absorption due to the fact that increases the solubility of phytic acid under low pH values. However, it is contradictory to the findings in the present study because boron supplementation linearly reduced the plasma phosphorus concentrations. Moreover, the impact of boron supplementation on phosphorus digestibility and plasma concentrations is inconsistent in the previous studies as reported by Mızrak et al [26]. Nevertheless, it has been found that increased boron supplementation to pigs increased phosphorous level in bone whereas calcium level was not impacted [2]. Therefore, boron may be efficiently involving in the deposition of phosphorous in bone from plasma rather than calcium. This concept was also suggested in Green [25].
The impact of boron supplementation on plasma cholesterol and triglycerides in pigs was reported in previous studies [2,3]. The outcomes of the present study are lining with the previous studies where boron supplementation increase plasma cholesterol level [2] and reduced plasma triglyceride levels [27]. However, contradictory results were reported in Armstrong et al [28] such that boron supplementation increased plasma triglyceride levels. It has been reported that boron impact on cholesterol is indirect because boron impact on plasma thyroxine level regulates the cholesterol level as thyroxin has an inverse relationship with plasma cholesterol [29, 30]. The mechanism behind the impact of boron on plasma triglyceride levels remains unclear and further researches needed to confirm suitable explanations. In the small intestine, fats in vegetable oil hydrolyzed to monoglycerides and free fatty acids and absorbed into epithelial cells and resynthesized to triglyceride before entering the systemic plasma [31]. Therefore, reduction of feed intake may be a possible reason for low plasma triglyceride levels in pigs offered 10 mg/kg boron diets.
Diarrhea in pigs mainly caused by the growth of putrefactive microorganisms in the hindgut that is accelerated by the accumulation of undigested protein. The quadratic effect of boron supplementation indicates that 5 mg/kg decreases the diarrhea index in the present study regardless of sanitary conditions for 1 to 14 days. Therefore, the reduction of incidence of diarrhea in pigs offered 5 mg/kg may be caused by improved crude protein digestibility in the present study.


The effect of boron supplementation as a form of boric acid on ADG and G:F ratio in pigs independent to the sanitary conditions such that 5 mg/kg of boron supplementation supported optimum ADG and G:F whereas ADFI was dependent on sanitary conations for 1 to 14 days post-wean. The advantageous effect of 5 mg/kg boron on crude protein, energy and DM digestibility may be the reasons for improved growth performance and low diarrhea index in weaner pigs. Furthermore, poor nutrient digestibility, low plasma triglyceride levels and reduced feed intake reflected the supplementation of 10 mg/kg boron to diets may cause a nutrient deficiency in weaner pigs.



We certify that there is no conflict of interest with any financial organization regarding the material discussed in the manuscript.


The study was supported by the National Institute of Animal Science (PJ016214).

Table 1
Ingredient and nutrient composition of the experimental diet (as-fed basis)
Ingredients (%)
 Corn 34.89
 Wheat 12.00
 Soybean meal, 44% 28.00
 Fish meal 5.00
 Dried whey 15.00
 Vegetable oil 3.10
 Limestone 0.80
 Monocalcium phosphate 0.36
 Vitamin-mineral premix1) 0.40
 L-Lys-HCl, 78.8% 0.28
 DL-Methionine 0.11
 Threonine 0.06
Calculated chemical composition
 Metabolizable energy (kcal/kg) 3,400
 Calcium (%) 0.85
 Methionine+cysteine (%) 0.84
 Lysine (%) 1.50
Analyzed composition
 Gross energy (kcal/kg) 3,900
 Crude protein (%) 22.32

1) Provided the following nutrients (per kg of air-dry diet): vitamin A, 7,000 IU; vitamin D3, 1,400 IU; vitamin E 20, mg; vitamin K, 1 mg; vitamin B1 (thiamine), 1 mg; vitamin B2 (riboflavin), 3 mg; vitamin B6 (pyridoxine), 1.5 mg; vitamin B12 (cobalamin), 15 μg; calcium pantothenate, 10.7 mg; folic acid, 0.2 mg; niacin, 12 mg; biotin, 30 μg; Co, 0.2 mg (as cobalt sulphate); Cu, 10 mg (as copper sulphate); iodine, 0.5 mg (as potassium iodine); iron, 60 mg (as ferrous sulphate); Mn, 40 mg (as manganous oxide); Se, 0.3 mg (as sodium selenite); Zn, 100 mg (as zinc oxide).

Table 2
Effects of environment and boron levels (mg/kg) on average daily gain (ADG, g/pig/d), average feed intake (ADFI, g/pig/d) and gain-to-feed ratio (G:F, g/g) in weaned pigs from day 1 to 14 post-weaning
Environment Boron levels Day 1–7 Day 8–14 Day 1–14

Good sanitary 0 230 158 0.69 530 545 1.03 380 351 0.93
5 368 275 0.74 652 660 1.01 510 467 0.91
10 178 182 1.01 312 203 0.60 245 192 0.74
Poor sanitary 0 271 195 0.69 613 548 0.86 442 371 0.83
5 233 152 0.46 556 661 1.17 394 407 0.99
10 180 174 1.05 398 299 0.78 289 237 0.87
SEM 32.2 41.0 0.134 54.6 67.4 0.083 39.1 44.5 0.056
--------------------------------------------------- Mean value of main effects ----------------------------------------------------------
 Good sanitary 258 205 0.86 500 473 0.88 379 342 0.87
 Poor sanitary 228 174 0.70 521 500 0.94 375 334 0.89
Boron levels
 0 250 176 0.73 573 549 0.95 412 366 0.89
 5 300 214 0.63 605 662 1.09 452 439 0.96
 10 179 178 0.99 353 248 0.69 266 210 0.79
------------------------------------------------------ Significance of main effects ----------------------------------------------------------
Environment 0.275 0.086 0.078 0.652 0.637 0.434 0.895 0.831 0.694
Boron supplementation
 Linear 0.036 0.813 0.016 <0.001 <0.001 0.004 0.001 0.001 0.078
 Quadratic 0.005 0.245 0.016 0.006 <0.001 0.001 0.002 <0.001 0.014
------------------------------------ Significance of interaction effects from orthogonal contrast ----------------------------
 Linear 0.538 0.570 0.868 0.976 0.486 0.049 0.817 0.781 0.051
 Quadratic 0.009 0.051 0.205 0.067 0.686 0.268 0.019 0.245 0.537

Each simple effect mean represents 6 replicate pens.

SEM, standered error of mean.

Table 3
Effects of environment and boron levels (mg/kg) on ileal morphology in weaned pigs on day 7 and 14
Environment Boron levels Day 7 Day 14

Villus height (V, μm) Crypt depth (C, μm) V:C ratio Villus height (V, μm) Crypt depth (C, μm) V:C ratio
Good sanitary 0 616 247 2.51 736 319 2.38
5 611 272 2.27 699 288 2.51
10 588 265 2.16 798 340 2.37
Poor sanitary 0 234 85 2.82 286 50 4.77
5 181 75 2.42 266 53 3.43
10 209 84 2.61 281 31 3.33
SEM 18.1 10.0 0.173 32.5 12.8 0.456
------------------------------------------------------ Mean value of main effects ------------------------------------------------------
 Good sanitary 625 250 2.26 763 309 2.49
 Poor sanitary 188 97 2.67 259 87 3.78
Boron levels
 0 424 167 2.65 510 196 3.60
 5 400 173 2.37 486 183 2.95
 10 396 180 2.39 537 214 2.84
------------------------------------------------------ Significance of main effects -------------------------------------------------------
Sanitary condition <0.001 <0.001 0.008 <0.001 <0.001 0.002
 Linear 0.130 0.186 0.126 0.422 0.187 0.106
 Quadratic 0.528 0.885 0.307 0.206 0.055 0.496
------------------------------- Significance of interaction effects from orthogonal contrast -----------------------------
 Linear 0.938 0.138 0.687 0.310 0.848 0.122
 Quadratic 0.124 0.286 0.437 0.376 0.074 0.351

Each simple effect mean represents 6 replicate pens.

SEM, standered error of mean.

Table 4
Effects of environment and boron levels (mg/kg) on apparent total tract digestibility (%) of nutrients in weaned pigs on days 7 and 14
Environment Boron levels Day 7 Day 14

Dry matter Crude protein Energy Dry matter Crude protein Energy
Good sanitary 0 92.4 89.6 90.2 92.9 88.3 91.6
5 94.6 91.8 93.9 93.2 91.7 92.5
10 88.6 80.5 86.8 91.0 85.4 90.3
Poor sanitary 0 88.9 75.4 87.2 87.4 79.2 84.9
5 91.5 86.8 90.3 92.7 89.8 91.6
10 82.3 74.9 81.4 87.5 80.6 86.7
SEM 0.83 1.73 0.91 1.63 2.88 1.33
--------------------------------------------------- Mean value of main effects ---------------------------------------------------------
 Good sanitary 91.9 87.3 90.3 92.4 88.4 91.4
 Poor sanitary 87.6 79.0 86.3 89.2 83.3 87.58
Boron levels
 0 90.6 82.5 88.7 90.1 83.6 88.1
 5 93.1 89.3 92.1 93.0 90.9 92.1
 10 85.4 777 84.1 89.3 83.0 88.5
----------------------------------------------------------- Significance of main effects -------------------------------------------------
Environment 0.001 <0.001 <0.001 0.018 0.024 0.025
Boron supplementations
 Linear <0.001 0.010 <0.001 0.595 0.840 0.834
 Quadratic <0.001 <0.001 <0.001 0.018 0.002 0.029
---------------------------- Significance of interaction effects from orthogonal contrast ---------------------------------
 Linear 0.771 0.013 0.723 0.132 0.230 0.149
 Quadratic 0.051 0.205 0.190 0.869 0.888 0.968

Each simple effect mean represents 6 replicate pens.

SEM, standered error of mean.

Table 5
Effects of environment and boron levels (mg/kg) on pro-inflammatory cytokines of gut mucosa in weaned pigs on days 7 and 14
Environment Boron levels Day 7 Day 14

IL-1β (pg/mL) TNF-α (pg/mL) IL-1β (pg/mL) TNF-α (pg/mL)
Good sanitary 0 23.6 1.2 65.5 1.8
5 9.6 3.3 11.4 1.7
10 24.6 0.7 56.3 1.2
Poor sanitary 0 14.1 31.7 15.7 2.3
5 69.0 2.6 44.7 1.9
10 43.1 1.6 26.8 2.9
SEM 27.11 16.93 31.16 1.13
---------------------------------------------------- Mean value of main effects ----------------------------------------------
 Good sanitary 19.3 1.7 44.4 1.6
 Poor sanitary 42.1 11.9 29.1 2.3
Boron levels
 0 18.9 16.4 40.6 2.1
 5 32.3 3.0 28.0 1.8
 10 33.9 1.1 41.5 2.0
------------------------------------------------------ Significance of main effects ---------------------------------------------
Environment 0.319 0.473 0.554 0.413
Boron supplementations
 Linear 0.597 0.379 0.379 0.984
 Quadratic 0.581 0.702 0.702 0.812
----------------------- Significance of interaction effects from orthogonal contrast -----------------------------
 Linear 0.622 0.395 0.749 0.604
 Quadratic 0.249 0.588 0.193 0.648

Each simple effect mean represents 6 replicate pens.

IL-1β, interleukin-1β; TNF-α, tumor necrosis factor-α; SEM, standered error of mean.

Table 6
Effects of environment and boron levels (mg/kg) on nutrient composition in systemic plasma of weaned pigs on days 7 and 14
Environment Boron levels Day 7 Day 14

Calcium (mg/dL) Phosphorous (mg/dL) Cholesterol (mg/dL) Triglyceride (mg/dL) Calcium (mg/dL) Phosphorous (mg/dL) Cholesterol (mg/dL) Triglyceride (mg/dL)
Good sanitary 0 10.5 7.4 59.1 78.2 12.2 7.8 68.1 114.3
5 10.7 7.2 60.8 43.1 12.4 7.2 68.2 79.4
10 11.0 6.7 62.6 32.6 12.9 7.2 78.3 74.1
Poor sanitary 0 10.2 7.1 79.2 59.1 12.2 10.2 68.9 98.7
5 10.7 6.2 53.7 49.4 12.7 9.3 68.3 70.7
10 10.7 5.5 61.1 35.1 13.0 9.0 84.4 65.2
SEM 0.21 0.28 4.23 7.83 0.25 0.48 6.18 12.41
-------------------------------------------------------------------- Mean value of main effects ------------------------------------------------------
 Good sanitary 10.8 7.1 60.9 51.3 12.4 7.4 71.9 89.3
 Poor sanitary 10.5 6.3 64.7 47.9 12.6 9.5 73. 78.2
Boron levels
 0 10.4 7.3 69.2 69.0 12.2 9.0 68.0 106.5
 5 10.7 6.7 57.3 46.2 12.5 8.3 68.8 75.0
 10 10.9 6.1 61.9 33.8 13.0 8.1 81.3 69.7
-------------------------------------------------------------------- Significance of main effects ------------------------------------------------------
Environment 0.214 0.001 0.276 0.595 0.338 <0.001 0.741 0.282
Boron supplementations
 Linear 0.026 <0.001 0.091 <0.001 0.004 0.070 0.037 0.005
 Quadratic 0.658 0.898 0.030 0.465 0.827 0.486 0.277 0.231
----------------------------------------- Significance of interaction effects from orthogonal contrast ---------------------------------
 Linear 0.966 0.080 0.014 0.173 0.765 0.612 0.613 0.789
 Quadratic 0.560 0.753 0.031 0.287 0.420 0.988 0.722 0.868

Each simple effect mean represents 6 replicate pens.

SEM, standered error of mean.

Table 7
Effects of environment and boron levels (mg/kg) on diarrhea index in weaned pigs on days 7 and 14
Environment Boron levels Day 1–7 Day 8–14 Day 1–14
Good sanitary 0 3.57 17.88 10.70
5 0.00 3.58 1.78
10 3.57 13.10 10.73
Poor sanitary 0 7.15 14.28 10.73
5 0.00 3.57 1.78
10 21.43 21.43 21.43
SEM 4.339 6.792 4.071
--------------------------------------------------- Mean value of main effects ----------------------------------------------
 Good sanitary 2.38 13.10 7.73
 Poor sanitary 9.52 13.10 11.31
Boron levels
 0 5.36 16.08 10.71
 5 0.00 3.57 1.78
 10 12.50 19.64 16.08
------------------------------------------------------- Significance of main effects -------------------------------------------
Environment 0.050 0.999 0.288
Boron supplementation
 Linear 0.107 0.603 0.195
 Quadratic 0.022 0.020 0.002
------------------------------- Significance of interaction effects from orthogonal contrast --------------------
 Linear 0.107 0.600 0.197
 Quadratic 0.161 0.999 0.451

Each simple effect mean represents 6 replicate pens.

SEM, standered error of mean.


1. Nielsen FH. Boron in human and animal nutrition. Plant Soil 1997; 193:199–208. https://doi.org/10.1023/A:1004276311956
2. Armstrong TA, Spears JW, Crenshaw TD, Nielsen FH. Boron supplementation of a semipurified diet for weanling pigs improves feed efficiency and bone strength characteristics and alters plasma lipid metabolites. J Nutr 2000; 130:2575–81. https://doi.org/10.1093/jn/130.10.2575
crossref pmid
3. Armstrong TA, Spears JW. Effect of dietary boron on growth performance, calcium and phosphorus metabolism, and bone mechanical properties in growing barrows. J Anim Sci 2001; 79:3120–7. https://doi.org/10.2527/2001.79123120x
crossref pmid
4. Cheng J, Peng K, Jin E, et al. Effect of additional boron on tibias of African ostrich chicks. Biol Trace Elem Res 2011; 144:538–49. https://doi.org/10.1007/s12011-011-9024-y
crossref pmid
5. Armstrong TA, Flowers WL, Spears JW, Nielsent FH. Long-term effects of boron supplementation on reproductive characteristics and bone mechanical properties in gilts. J Anim Sci 2002; 80:154–61. https://doi.org/10.2527/2002.801154x
crossref pmid
6. Armstrong TA, Spears JW. Effect of boron supplementation of pig diets on the production of tumor necrosis factor-α and interferon-γ. J Anim Sci 2003; 81:2552–61. https://doi.org/10.2527/2003.81102552x
crossref pmid
7. Białek M, Czauderna M, Krajewska KA, Przybylski W. Selected physiological effects of boron compounds for animals and humans. A review. J Anim Feed Sci 2019; 28:307–20. https://doi.org/10.22358/jafs/114546/2019
8. Shin TK, Yi YJ, Kim JC, et al. Reducing the dietary omega-6 to omega-3 polyunsaturated fatty acid ratio attenuated inflammatory indices and sustained epithelial tight junction integrity in weaner pigs housed in a poor sanitation condition. Anim Feed Sci Technol 2017; 234:312–20. https://doi.org/10.1016/j.anifeedsci.2017.04.022
9. Humphrey BD, Klasing KC. Modulation of nutrient metabolism and homeostasis by the immune system. World Poult Sci J 2004; 60:90–100. https://doi.org/10.1079/WPS20037
10. Kim JC, Hansen CF, Mullan BP, Pluske JR. Nutrition and pathology of weaner pigs: nutritional strategies to support barrier function in the gastrointestinal tract. Feed Sci Technol 2012; 173:3–16. https://doi.org/10.1016/j.anifeedsci.2011.12.022
11. National Research Council. Nutrient requirements of swine. 11th edWashington, DC, USA: National Academy Press; 2012.

12. Heo JM, Kim JC, Hansen CF, Mullan BP, Hampson DJ, Pluske JR. Feeding a diet with decreased protein content reduces indices of protein fermentation and the incidence of postweaning diarrhea in weaned pigs challenged with an enterotoxigenic strain of Escherichia coli. J Anim Sci 2009; 87:2833–43. https://doi.org/10.2527/jas.2008-1274
crossref pmid
13. Marquardt RR, Jin LZ, Kim JW, Fang L, Frohlich AA, Baidoo SK. Passive protective effect of egg-yolk antibodies against enterotoxigenic Escherichia coli K88+ infection in neonatal and early-weaned piglets. FEMS Immunol Med Microbiol 1999; 23:283–8. https://doi.org/10.1111/j.1574-695X.1999.tb01249.x
crossref pmid
14. Heo JM, Kim JC, Hansen CF, Mullan BP, Hampson DJ, Pluske JR. Feeding a diet with a decreased protein content reduces both nitrogen content in the gastrointestinal tract and post-weaning diarrhoea, but does not affect apparent nitrogen digestibility in weaner pigs challenged with an enterotoxigenic strain of Escherichia coli. Anim Feed Sci Technol 2010; 160:148–59. https://doi.org/10.1016/j.anifeedsci.2010.07.005
15. Pluske JR, Williams IH, Aherne FX. Maintenance of villous height and crypt depth in piglets by providing continuous nutrition after weaning. Anim Sci 1996; 62:131–44. https://doi.org/10.1017/S1357729800014417
16. Association of Official Analytical Chemists. In : Official methods of analysis: Changes in Official Methods of Analysis Made at the Annual Meeting; Supplement 15AOAC; 1990.

17. Piñeiro C, Piñeiro M, Morales J, et al. Pig-MAP and haptoglobin concentration reference values in swine from commercial farms. Vet J 2009; 179:78–84. https://doi.org/10.1016/j.tvjl.2007.08.010
crossref pmid
18. Stein HH, Fuller MF, Moughan PJ, et al. Definition of apparent, true, and standardized ileal digestibility of amino acids in pigs. Livest Sci 2007; 109:282–5. https://doi.org/10.1016/j.livsci.2007.01.019
19. Zhao J, Harper AF, Estienne MJ, Webb KE, McElroy AP, Denbow DM. Growth performance and intestinal morphology responses in early weaned pigs to supplementation of antibiotic-free diets with an organic copper complex and spray-dried plasma protein in sanitary and nonsanitary environments. J Anim Sci 2007; 85:1302–10. https://doi.org/10.2527/jas.2006-434
crossref pmid
20. Ao Z, Kocher A, Choct M. Effects of dietary additives and early feeding on performance, gut development and immune status of broiler chickens challenged with Clostridium perfringens. Asian-Australas J Anim Sci 2012; 25:541–51. https://doi.org/10.5713/ajas.2011.11378
crossref pmid pmc
21. Goldbach HE, Wimmer MA. Boron in plants and animals: is there a role beyond cell-wall structure? J Plant Nutr Soil Sci 2007; 170:39–48. https://doi.org/10.1002/jpln.200625161
22. Suiryanrayna MV, Ramana JV. A review of the effects of dietary organic acids fed to swine. J Anim Sci Biotechnol 2015; 6:45 https://doi.org/10.1186/s40104-015-0042-z
crossref pmid pmc
23. Kabay N, Bryjak M. Boron removal from seawater using reverse osmosis integrated processes. Kabay N, Bryjak M, Hilal N, editorsBoron separation processes. Amsterdam, Netherland: Elsevier; 2015. p. 219–35. https://doi.org/10.1019/B978-0-444-63454-2.00009-5
24. Bhasker TV, Gowda NKS, Mondal S, et al. Boron influences immune and antioxidant responses by modulating hepatic superoxide dismutase activity under calcium deficit abiotic stress in Wistar rats. J Trace Elem Med Biol 2016; 36:73–9. https://doi.org/10.1016/j.jtemb.2016.04.007
crossref pmid
25. Green D. Effects of boron on selected aspects of swine health related to calcium and phosphorus metabolism. Carbondale, IL, USA: Southern Illinois University; 2020. Available from: https://opensiuc.lib.siu.edu/gs_rp

26. Mızrak C, Yenice E, Can M, Yıldırım U, Atik Z. Effects of dietary boron on performance, egg production, egg quality and some bone parameters in layer hens. S Afr J Anim Sci 40:257–64. https://doi.org/10.4314/sajas.v40i3.10
27. Kabu M, Uyarlar C, Zarczynska K, Milewska W, Sobiech P. The role of boron in animal health. J Elem 2015; 20:535–41. https://doi.org/10.5601/jelem.2014.19.3.706
28. Armstrong TA, Spears JW, Engle TE, Wright CL. Effect of dietary boron on bone characteristics and plasma parameters in young pigs. Roussel AM, Anderson RA, Favier AE, editorsTrace Elements in Man and Animals. 10:New York, NY, USA: Springer; 2002. p. 1067–69. https://doi.org/10.1007/0-306-47466-2_326
29. Field FJ, Albright E, Mathur SN. Effect of dietary cholesterol on biliary cholesterol content and bile flow in the hypothyroid rat. Gastroenterology 1986; 91:297–304. https://doi.org/10.1016/0016-5085(86)90560-3
crossref pmid
30. Eder K, Stangl GI. Plasma thyroxine and cholesterol concentrations of miniature pigs are influenced by thermally oxidized dietary lipids. J Nutr 2000; 130:116–21. https://doi.org/10.1093/jn/130.1.116
crossref pmid
31. Lairon D. Digestion and absorption of lipids. McClements J, Decker EA, editorsDesigning functional foods. Woodhead Publishing; 2009. p. 68–93. https://doi.org/10.1533/9781845696603.1.66

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