Table 3. Amelioration of heat stress in livestock using different nutritional interventions

Reference Animal HS condition Amelioration Major effects in comparison with HS versus ameliorating agents under HS
[145] Hubbard chickens 35°C, 75% RH, 8 h/day from 21 to 42 days MOS (0.5% in diet) and/or probiotics (0.1% in diet) ● MOS increased crypt depth in HS birds.● Villus height and surface area increased in ileum due to MOS and probiotic supplementation under HS.● Villus width and crypt depth increased in HS-probiotic group.● MOS, probiotic and MOS+probiotic enhanced activity of goblet cells under HS.
[84] Broiler chickens at 36 days of age 36°C for 0 to 10 h for one day Enhancer (L-glutamine injected intraperitoneally, 0.75 mg/kg BW) vs. inhibitor (quercetin injected intraperitoneally, 5 mg/kg of BW), 1 day before HS ● Increased GPx and SOD activity, T-AOC and HSP70 in jejunal mucosa by enhancer than by inhibitor.● LDH in jejunal mucosa was lower in enhancer group than inhibitor group.● LDH activity rapidly increased in jejunum during the first 2 h of HS.● Increased serum corticosterone in inhibitor group compared with enhancer group.
[85] Male broiler chickens 36°C for 0 to 10 h for one day Intraperitoneally injected with L-glutamine (enhancer, 0.75 mg/kg of BW) or quercetin (inhibitor, 5 mg/kg of BW) for one day before HS ● No effect on villus height, crypt depth (except decrease in 5 h), or VCR (except increase in 10 h) HSP70 mRNA expression in enhancer group was higher than inhibitor group in jejunum.● Trypsin, amylase, and lipase activities in enhancer group were higher than in inhibitor group.
[141] Ross-708 of mixed sex chickens 35°C from 1 to 42 days 0.5% MOS or 0.1% PM ● MOS and PM increased BW gain and ADFI, and lowered FCR compared with HS-control group.● Probiotic increased villus width and surface area compared with HS-control treatment.● MOS and PM partially alleviated changes of intestinal microstructures damaged by HS.
[146] Wenchang male chickens 40°C, 52.4% RH for 3 h 0.2 mL gamma-amino butyric acid solution (50 mg/kg of BW) daily for 35 days ● Increased GPx, SOD, catalase, T-AOC content in intestinal mucosa.● Decreased MDA content in jejunal and ileal mucosa.
[147] 21-day-old Ross male broilers 33°C from 08:00 to 06:00 from 21 to 42 days COS at 1.5 g/kg diet ● No effect on BW gain, feed intake, and FCR.● Increased number of Lactobacillus, decreased E. coli, and had no effect on Bifidobacterium and Clostridium counts.● Increased villus height and VCR, no effect on crypt depth.● Reduced jejunal FITC-d permeability.
[142] Crossbred gilts (43 kg BW) 36°C, 50% RH for either 1 or 7 days 100 (Zn220) and 200 (Zn320) mg/kg diet of Zn as Zn-amino acid complexes ● BW and feed intake were not affected.● Improved Ileal and colonic TER in Zn220-fed pigs● Ileal and colonic FITC-d permeability increased, and colonic FITC-LPS permeability tended to increase due to HS from days 1 to 7; but Zn supplementation has no effect.● Overall, no effects on ileal glucose, lysine, methionine, and glutamine transport.● Glucose transport linearly decreased with increasing levels of Zn-amino acid on day 1.● No differences on villi height or crypt depth.
[143] Crossbred gilts (39 kg BW) 32°C, 26% RH for 24 h WP (80% 98% and 100%) ● BW, BW gain, and feed intake were not affected by whey protein.● Plasma L-lactate and D-lactate levels increased and tended to increase with HS.● After 24 h of HS, 100% WP-fed pigs had lower plasma D-lactate relative to control-fed pigs.● Decreased Ileal TER (37%) in 80% WP-fed pigs.● Ileal TER decreased in HS whey protein groups.
[148] Hubbard male broiler chicks 35°C and 64% RH from 09:00 to 14:00 and 21°C from 14:00 to 09:00 Bacillus subtilis (1 g/kg diet; 2.3 × 108 CFU/g of B. subtilis spores) ● Increased BW, ADG and feed efficiency, but no effect on feed intake by B. subtilis.B. subtilis increased villus height, villus surface area, and absorptive epithelial cell area in duodenum and ileum.B. subtilis increased Lactobacillus and Bifidobacterium and decreased coliforms and Clostridium counts in intestine.
[131] Male Wenchang chickens 40.5°C, 52.4% RH for 2 h for 15 days GABA (50 mg/kg of BW) ● HS decreased activity of sucrase, maltase, alkaline phosphatase, contents of secretory IgA, glutathione, d-xylose, number of lymphocytes, and Na+-K+-ATPase activity of intestinal mucosa.● GABA ameliorated the above effects.
[7] 21-day-old Ross male broiler chickens 33°C for 10 h/day, from 22 to 42 days PM (B. licheniformis, B. subtilis, and Lactobacillus plantarum) at 1.5 g/kg diet ● Probiotic decreased FCR, but no effect on ADG and ADFI.● Probiotics had no effect on TER value and FITC-d permeability in jejunum compared with control diet.● Probiotics increased protein level of OCLN.● Probiotics increased villus height than the control diet.
[97] Male Wenchang chickens 40.5°C, 52.4% RH for 2 h for 15 days GABA (0.5%) fed with 0.2 mL ● GABA increased BW, but no effect on ADG, FCR and fed intake on day 15.● GABA enhanced villus height, crypt depth, mucosa thickness, intestinal wall thickness, and number of goblet cells in duodenum and ileum.
[135] Male Cobb 500 broilers birds 32°C–27°C vs. 37°C–33°C, decreased 2°C/week until reaching 33°C in third week; for 5 h/day from 29 to 42 day of age 10 g/kg diet of glutamic acid ● Glutamic acid increased villus height, crypt depth, and VCR.● Glutamic acid increased BW.
[149] Cobb 500 male broiler chickens 34°C, 65%–70% RH for 5 h/day from 29 to 42 days GSE 150, 300, 450 mg/kg of diet and vitamin C at 300 mg/kg of diet ● GSE or vitamin C did not affect lengths of duodenum, jejunum, ileum, large intestine and cecum.● GSE increased villus height, tended to increase crypt depth, but no effect on VCR and muscle layer thickness jejunum.● No effect of vitamin C on jejunum villus width, crypt depth, VCR, and muscle layer thickness.● GSE, but not vitamin C, lowered ileal coliforms and E. coli populations.
[99] Crossbred gilts (64 kg BW) 37°C and 40% RH for 12 h ZnAA ● No differences of BW loss and feed intake between HS-control and HS-ZnAA pigs.● ZnAA decreased BW and feed intake under HS.● ZnAA had no effect on FITC-d permeability, but increased ileal TER level under HS.● Ileal IL1-β decreased and plasma LBP increased in HS-ZnAA.● Colon MUC2 mRNA abundance elevated in HS-ZnAA AA pigs● Intestinal alkaline phosphatase gene abundance increased in HS-ZnAA.● No effect on ileal and colonic HIF1A protein expression.● No effect on ileal HSP70 protein expression but lowered HSP70 protein expression in colon of HS-ZnAA pigs.
[1] 15-day-old Ross broiler chickens 38°C–39°C for 8 h for 5 days GOS at 10 or 25 g/kg diet (6 days prior to HS) ● GOS prevented HS-induced upregulation of HSF3, HSP70, and HSP90 mRNA in jejunum.● Heat-induced effects on CDH1 mRNA expression and pan-cadherin protein expression in jejunum prevented by GOS.● HS-induced increase in mRNA expressions of CLDN5 and ZO1 dose-dependently alleviated in jejunum by GOS.● GOS (25 g/kg) prevented heat-induced gene TLR-4 induction in jejunum.● Up-regulation of IL6 and IL8 mRNA expressions in jejunum and ileum by HS was prevented by GOS.
[71] 28 day old female Xuefeng black-boned chickens 37°C for 8 h/day for 15 days 200, 400, or 600 mg/kg diet of resveratrol for 15 days ● Resveratrol improved villus morphology, increased goblet cell and lymphocyte numbers at 400 mg/kg.● Resveratrol reduced HSP70, HSP90, and NF-κB protein levels in jejunal villi after 15 days of HS and increased EGF level.
[130] 21 day old Hubbard male broiler chickens 32°C; 64% RH 0.5 g butyric acid/kg of feed ● Butyric acid increased BW and ADG in HS birds.● Butyric acid increased recovery of villus height, villus surface area, relative intestinal weight, and absorptive epithelial cell area in duodenum of HS-birds.● Butyric acid increased intestinal Lactobacillus and Bifidobacterium populations in HS-birds.
[94] Female growing pigs 35°C 8 h/day, 35% RH for 2 days 0.2–1.0 mg/kg of Se and 17–200 IU/ kg of vitamin E for 14 days ● Linearly increased TER value.● Decreased FITC-d permeability quadratically.● Increased GPx2 mRNA abundance.● Se and vitamin E did not affect HSP70, HIF1A, IL8, or TNFA mRNA levels.
[30] Hy-Line Brown laying hens (aged 40 weeks) 33°C and 60%–70% RH, for 20 days PM (Enterococcus faecium and B. subtilis) ● Probiotic increased egg production rate, feed intake, and egg weight under HS.● Probiotic improved eggshell strength, eggshell thickness, and albumen height.● Probiotics increased VCR in ileum (10 days and 20 days) and cecum (20 days) in HS-hens.● Probiotic upregulated expression levels of OCLN, ZO1, and JAM-A in ileum and cecum.● Probiotics reduced mRNA level of HSP70.
[93] Cobb male chickens 36°C from 08:00 to 18:00 and 26°C from 18:00 to 08:00 from 8 to 35 days NAC (1 g/kg in diet) ● NAC increased ADFI, ADG, and reduced FCR.● NAC increased VCR.● NAC increased ATP level in jejunal mucosa.● NAC inhibited the increase of jejunal AMP under HS.● NAC decreased MDA concentration and mRNA levels of HSP70, AMP-activated protein kinase, and heme-oxigenase in intestine.● NAC elevated catalase and trypsine activity in jejunum under HS.
[66] Lohmann layer cockerels TN (21°C, 62% RH) vs. HS (35°C, 64% RH from 09:00 to 13:00 and 21°C from 13:00 to 09:00) for 30 days Butyrate at 0.35 g/kg of diet ● Butyrate increased villus height, surface area, and absorptive epithelial cell area in all intestinal parts under HS.● No effect of butyrate on crypt depth in duodenum and jejunum under HS, but it was greater in ileum.● Butyrate alleviated villi damages and epithelial cell damage in all intestinal sections.● Butyrate increased intestinal and cecal Lactobacillus and Bifidobacterium numbers and decreased Clostridium and coliforms numbers.
[86] 21-day-old Cobb male broiler chickens 33°C, 70% RH for 10 h for 21 days 400 mg/kg diet of resveratrol ● Resveratrol increased final BW but no effect on ADG and ADFI.● Resveratrol increased villus height, VCR and goblet cell numbers, and lowered crypt depth.● Resveratrol increased numbers of Lactobacillus and Bifidobacterium and lowered the numbers of Escherichia coli colonization.● Resveratrol increased intestinal mucosal CLDN1, OCLN, CDH1, and MUC2 mRNA levels.● Resveratrol decreased serum D-lactic acid and FITC-d concentrations in HS birds.
[62] Crossbred gilts (50 kg BW) 27°C–30°C, 35% humidity for 7 days Zn at 120 mg/kg diet (60 mg as Zn-sulfate + 60 mg as Zn-amino acid) ● No effect on ADFI, ADG, and final BW, but FCR tended to decrease.● No effect on TNFA and LPS-binding protein in serum.
[144] 21-day-old Arbor Acres broiler chickens 36°C for 10 h/day for 20 days 5 and 10 g/kg diet of glutamine ● Glutamine increased villus height in HS birds.● Glutamine decreased D-lactate and diamine oxidase activity in HS birds.● Glutamine mediated secretion of cytokines (TNFA and IL10), increased ZO1, CLDN1, and OCLN mRNA levels in HS birds.
[76] Male Arbor Acres plus broiler chickens Control (20°C) vs. HS (32°C–33°C 8 h/day) for 4–42 days MOS at 250 mg/kg diet ● MOS increased ADG, ADFI, and FCR.● MOS decreased mucosal MDA content in jejunum at 42 days under HS.● MOS increased jejunal and ileal villus height and VCR.● MOS increased mRNA abundances of OCLN, and ileal MUC2 and ZO1 in jejunum and ileum, and ileal CLDN3 genes expressions.
[138] 28-day old male Ross-308 broiler chickens 33°C and 40%–55% humidity for 8 h/day for 10 days 1 g/kg of betaine in finisher diet ● Betaine increased ileal villus height and TER.● Betaine decreased Evans blue dye concentration in jejunum and ileum.
[134] Male Arbor Acres broilers from 28 to 42 days TN (23°C) or subjected to cyclic HS (28°C–35°C–28°C for 12 h daily) Probiotic (Lactobacillus acidophilus, Lactobacillus plantarum, and Enterococcus faecalis) at a dose of 1.5 × 108 CFU/kg ● Probiotics had no effect on growth performance except for increased ADFI on days 22–42.● Probiotic increased villus height and VCR in duodenum and jejunum of HS birds on day 42.● Probiotic decreased serum D-lactic acid concentration on day 28.● Probiotic reduced serum TNFA and IL6, but increased IL10 and TGFB1 in HS birds.
[139] Male Cobb50 broiler chickens TN (26°C) vs. HS (34°C for 8 h daily for 21 days) 112.5 mg ginseng extract /kg feed ● Ginseng increased feed intake, BW and FCR.● Ginseng decreased HSPA1A, HSPD and HSPB1 gene expression.● Ginseng upregulated CLDN3, OCLN and CLDN1 (Caco 2).
HS, heat stress; RH, relative humidity; MOS, mannan-oligosaccharides; PM, probiotic mixture; BW, body weight; GPx, glutathione peroxidase; SOD, superoxide dismutase; T-AOC, total antioxidant capacity; HSP, heat shock protein; LDH, lactate dehydrogenase; VCR, villus height to crypt depth ratio; ADFI, average daily feed intake; FCR, feed conversion ratio; PM, probiotic mixture; MDA, malondialdehyde; FITC-d, fluorescein isothiocyanate labeled dextran; TER, transepithelial electrical resistance; WP, whey protein; GABA, γ-aminobutyric acid; GSE, grape seed extract; ZnAA, zinc amino acid complex; IL, interleukin; LBP, lipopolysaccharide binding protein; MUC, mucin; GOS, galacto-oligosaccharides; CDH1, E-cadherin; CLDN, claudin; ZO1, zonula occludens 1; TLR, toll-like receptor; NF-κB, nuclear factor kappa B; EGF, epidermal growth factor; HIF1A, hypoxia-induced factor-1α; TNFA, tumour necrosis factor α; JAM-A, junctional adhesion molecule A; NAC, N-acetylcysteine; AMP, adenosine monophosphate; TN, Thermoneutral; OCLN, occludin.