Table 1.
Condition, animal model | Probiotic, dose, vehicle, time | Offspring outcomes (compared to control) | Effect on gut microbiota in dams and offspring | Ref | |
---|---|---|---|---|---|
Before pregnancy | |||||
Metabolic disorders | |||||
SPF C57BL/6 J mice receiving a HFD or not since preconception ± probiotics Dams n = 9–10/group Offspring n = 27–30/group (male and female) |
Bifidobacterium breve DM8310, Lactobacillus acidophilus DM8302, Lacticaseibacillus casei DM8121, Streptococcus thermophilus DM8309 2 × 109 CFU/day by gavage 6 weeks before mating–PND21 |
↓ body weight until PND42 PND21: ↓ total cholesterol, LDL Male: ↓ HDL PND42: ↓ total cholesterol, LDL Female: ↓ glucose and insulin. ↑ HDL compared to control diet |
Dams ↓ Bacteroidetes, ↑ Bacteroidetes S24-7, Allobaculum, Sutterella Offspring PND21: Ameliorate HFD-induced dysbiosis (Prevotella, Bacteroides, Bacteroidales S24-7). Female: ↑ ⍺-diversity PND42: No effect on β-diversity. Female: Ameliorate dysbiosis (↑ Bacteroidaceae, Lachnospiraceae, Sutterella; ↓ Lactobacillus, Prevotella, Helicobacter, Rikenellaceae, Parabacteroides) |
[93] | |
Parasite infection | |||||
Swiss mice ± probiotics, inoculated with Toxocara canis on GD14 Dams n = 8/group Offspring n = 62–65/group (male and female) |
Saccharomyces boulardii CNCM I-745 1 × 107 CFU/day, added in feed 2 weeks before mating–PND21 |
PND21: ↓ 42% in the number of larvae transmitted, ↓ 50% larvae found in the brain |
Not assessed | [110] | |
During pregnancy | |||||
Wistar rats ± probiotics Dams n = 6/group Offspring n = 48/group (male and female) |
Limosilactobacillus fermentum CECT5716 1 × 1010 CFU/day by gavage GD1–PND14 |
No effect on body weight or BMI PND14: Intestine: ↑ IgA Plasma: ↑ IgG2a, ↓ IgG2c, ↑ Th2-type Ig, no change cytokines; ↓ in palmitic acid and total saturated fatty acids, ↑ plasma linoleic acid, eicosadienoic acid, 17:0 and 22:0 carbons fatty acids |
Dams & Offspring PND14: No effect on cecal microbiota composition, diversity, or richness PND21: Strain detected in all the dams cecal content |
[58] | |
SPF C57BL/6 mice ± probiotics Dams n = 10/group Offspring n = 8–10/group/time point (male and female) |
Akkermansia muciniphila MucT (ATCC BAA-835) 1 × 108–109 CFU/day by gavage GD1–3 days before birth; PND3–PND21 |
Until PND42: No effect on body weight or intestinal tissue development |
Dams No effect during pregnancy PND21: ↑ A. muciniphila, Ruminococcus_1, ↓ Coriobacteriaceae UCG-002, Lachnospiraceae UCG-001, Ruminiclostridium Offspring No effect on A. muciniphila abundance or ⍺-diversity PND42: Change in β-diversity, ↓ Dubosiella, Lachnospiraceae_UCG_001, Parasutterella, ↑ Gordonibacter, Lachnospiraceae_UCG_006, Prevotellaceae_UCG_001; ↑ the D-glutamine and D-glutamate microbial metabolism pathway |
[98] | |
Bama mini-sows ± probiotics or synbiotics Dams n = 16/group Offspring n = 8/group (male and female) |
Mix: Lactiplantibacillus plantarum B90, Saccharomyces cerevisiae P11 Dose undisclosed, added in feed Synbiotic: probiotic mix + xylo-oligosaccharides (500 g/t feed) GD1–PND28 |
PND58 Probiotics: ↑ CAT, GPx and SOD activities in plasma, jejunum, and colon; ↓ malondialdehyde and H2O2 concentrations in plasma Synbiotics: ↑ CAT and SOD activities in plasma; ↑ glutathione, GPx activities and mRNA levels of antioxidant and mitochoindrial-related genes in jejunum; ↑ total antioxidant capacities in colon; ↓ colonic malondialdehyde |
Offspring PND58 Probiotics: ↑ Bacteroidetes relative abundance in the jejunum and Bifidobacterium in the jejunum and colon. No change of total bacteria, Bacteroides, Clostridium cluster IV, Lactobacillus in jejunum or colon Synbiotics: ↑ Firmicutes, Bacteroidetes, Bifidobacterium, Lactobacillus (jejunum) |
[69] | |
Bama mini-sows ± probiotics or synbiotics Dams n = 16/group Offspring n = 8/group (male and female) |
Mix: Lactiplantibacillus plantarum B90, Saccharomyces cerevisiae P11 Dose undisclosed, added in feed Synbiotic: probiotic mix + xylo-oligosaccharides (500 g/t feed) GD1–PND28 |
PND65 Probiotics: Improve immune response (Plasma: ↓ IL-2 and LPS concentrations, ↑ IgA; Jejunum: ↑ IL-10, interferon-α, sIgA; ileum: ↑ sIgA) ↑ jejunal villus height, villus height/crypt depth ratio Synbiotics: ↑ plasma IgA, ↑ jejunal IL-10, interferon-α, sIgA, ↑ ileal villus height |
Offspring PND65: Effect on β-diversity, no effect on ⍺-diversity Probiotics: ↑ relative abundance of Psychrobacter, Dialister, Oleomonas, and Facklamia (ileum) Synbiotics: ↑ Turicibacter, SMB53, Clostridium, Paracoccus, Thalassobius, Vibrio, Psychrobacter, and Blautia (jejunum); ↑Corynebacterium and Agrobacterium (ileum) |
[70] | |
White Rex rabbit ± low /middle/high dose probiotics Dams n = 20/group Offspring n = 30/group (male and female) |
Clostridium butyricum CCTCC AB: 2017089 1 × 103 CFU/g (low dose), 1 × 104 CFU/g (middle dose), 1 × 105 CFU/g (high dose) in feed GD1–PND35, PND35–PND63 (offspring) |
PND28: ↑ average daily weight gain PND63: ↑ α-amylase and chymotrypsin activity in SI; ↑ SI villi length and ↓ crypt depth; modulate SOD, GPx and CAT activity ↑ ZO-1, claudin, and occludin mRNA in SI High dose: ↓ IL-6, TNF-α and IFN-γ in ileum and colon; ↑ sIgA in duodenum. ↑ MyD88, TLR2, and TLR4 relative expression in SI |
Offspring: PND63 Dose dependent effects High and middle dose: ↑ total bacteria abundance, ↑ Firmicutes; ↑ Lactobacillus & Bifidobacterium; ↑ Clostridium cluster IV, Clostridium cluster XIVa, and Butyrivibrio fibrisolvens in small intestine ↑ Clostridium cluster XIVa in colon Low dose: ↑ Clostridium cluster IV, Clostridium cluster XIVa, Butyrivibrio fibrisolvens, and Lactobacillus in duodenum |
[99] | |
C57BL/6 J mice ± probiotics Dams Undisclosed Offspring n = 9–10/group (male) n = 10–12/group (female) |
Lactococcus lactis 5 × 105 CFU/ml in drinking water GD10.5–PND1 |
PND1: ↑ the density of cortical neurons (Tbr1-expressing neurons, Satb2-expressing neurons in male), ↑ the density of blood vessels in the cortical plate Females: ↑ mitotic neural progenitor cell numbers 10 weeks: ↓ Anxiety-like behaviors Females: ↓ Freezing time in cue associated learning |
Not assessed | [77] | |
SPF Sprague Dawley rats ± probiotics Dams n = 4–5/group Offspring n = 12–14/group/time point (male) n = 11–14/group/time point (female) |
Lactobacillus helveticus NS8 1 × 108 CFU/ml in drinking water GD13–GD22 |
↓ body weight from PND51 until PND100-105 Antianxiety effect: PND35-40: Elevated plus maze: ↑ proportion of time in the open arms; PND35-40 & PND100-105: Open field test: ↑ proportion of the time/entries spent in the center and the distance travelled in the center, ↓ proportion of the latency to reach the center from the peripheral zone |
Not assessed | [63] | |
C57BL/6 mice + probiotic or fixed probiotic Dams n = 9/group Offspring n = 3–4/group/time point (male) n = 3–4/group/time point (female) |
Lacticaseibacillus rhamnosus GG 1 × 108 CFU/day (dams) by gavage, 1 × 107 CFU/day (offspring) in feed GD18–birth, PND1–PND5 (offspring) |
No impact on body weight PND21: ↑ crypt depth, villus height (jejunum, ileum, colon); 8 months: ↑ goblet cells, ZO-1, Claudin-1, and Occludin mRNA expression; ↑ ZO-1 and Claudin-3 protein expression ↑ SOD 1 & 2 mRNA expression, GPx expression |
Offspring PND21: Strain detected in feces; no impact on ⍺-diversity, impact β-diversity; ↑ Akkermansia muciniphila and SCFA-producing bacteria (Ruminococcus, Coprococcus, Odoribacter, Faecalibaculum, and Lachnospiraceae bacterium A4) 8 months: Effect on β-diversity maintained ↑ Lactobacillus, Parasutterella, Bifidobacterium, Akkermansia muciniphila, ↓ Oscillibacter, Escherichia, Ruminococcus, Helicobacter, Alistipes timonensis |
[59] | |
C57BL/6 J mice + probiotic or fixed probiotic Dams Undisclosed Offspring n = 11–16/group (male) n = 10–13/group (female) |
Lacticaseibacillus rhamnosus GG 1 × 108 CFU/day (dams) by gavage, 1 × 107 CFU/day (offspring) in feed GD18–birth; PND1–PND5 (offspring) |
↑ BW at PND7, 14, 21, 28 PND21: ↑ Intestinal villus length, colonic crypt depth, cell proliferation and differentiation, tight junction proteins expression (ZO-1, claudin-3) 14 weeks (males): ↑ Fecal SCFA concentrations, ↓ anxiety-like behavior, ↑ colon epithelial growth factor receptor activation, serotonin transporter mRNA and protein levels; ↓ colon and serum 5-HT levels, ↑ brain-derived neurotrophic factor and ɣ-aminobutyric receptor |
Offspring PND21: No effect on ⍺-diversity, impact β-diversity, ↑Akkermansia, ↓ Helicobacter, Flavonifractor, Oscillibacter, Alistipes ↓ bacterial pathways (motility proteins, flagellar assembly and porphyrin and chlorophyll metabolism) 14 weeks (male): No effect on ⍺-diversity, impact β-diversity, ↑ Clostridium XIVa, Bifidobacterium ↓ Prevotella, Akkermansia, Bifidobacteria |
[62] | |
Landrace × Yorkshire sows ± probiotics; offspring ± synbiotics (split-plot design) Dams 14–15/group Offspring n = 4–5/pen, 18–19 pens/group (male and female) |
Bacillus subtilis C-3102 5 × 105 (gestation), 1 × 106 (lactation), 5 × 105 (offspring) CFU/g of feed Prebiotics (MOS, offspring) GD30–PND19; PND19–PND42 (offspring) |
No effect of nursery treatment on overall growth PND42: ↓ Weight in piglets born from probiotic supplemented sows |
Dams GD30, GD113, PND18: ↑ Fecal Bacillus subtilis C-3102 & total Bacillus sp. Offspring PND18: ↑ Fecal Bacillus subtilis C-3102 & total Bacillus sp. |
[111] | |
Great Dane Bitches ± synbiotics (started at GD35 or GD56) Dams n = 5/group Offspring n = 30–32/group (male and female) |
Mix: Enterococcus faecium DSM 10663, Lactobacillus acidophilus CECT 4529 3.36 × 108 and 1.03 × 1010 CFU/10 kg of BW, respectively, in tablets Prebiotics (FOS: 480 mg/10 kg, MOS: 48.6 mg/10 kg of BW) GD35 or 56–birth |
No effect on birth weight or litter parameters 9 weeks: ↓ Incidence of gastroenteritis 4-week supplementation had better results on first presentation of gastroenteritis |
Not assessed | [112] | |
Large White sows ± probiotics Dams n = 20/group Offspring n = undisclosed (male and female) |
Mixes: Bacillus subtilis A + Bacillus subtilis B (BS-A + B); Bacillus subtilis A + Bacillus licheniformis (BS-A + BL); or Bacillus subtilis B + Bacillus licheniformis (BS-B + BL) Total of 4 × 109 CFU/kg of feed GD85–PND21 |
PND21: ↑ Average weight and average daily gain at weaning BS-A + BL: ↑ number of weaned piglets |
Dams GD100-112: No effect on Bifidobacterium spp., Escherichia coli or total bacteria BS-B + BL: GD112: ↑ Clostridium cluster IV PND7: ↓ Firmicutes |
[60] | |
Landrace x Yorkshire sows ± probiotics Dams n = 16/group Offspring n = 12/dams (male and female) |
Bacillus subtilis PB6 4 × 108 CFU/kg of feed GD90–PND21 |
↑ litter size, ↓ BW PND14: ↓ Serum cortisol concentration PND21: ↑ Litter weights and litter weight gains |
Dams GD110: ↓ ⍺-diversity, ↑ Gemmatimonadetes, Acidobacteria, Ruminococcaceae_UCG-013 cc, ↓ Proteobacteria, Actinobacteria, Streptococcus relative abundance |
[61] | |
Pietrain sows ± probiotics Dams n = 10/group Offspring n = undisclosed (male and female, n = 5/group for hematological profile) |
Lactiplantibacillus plantarum CAM6 1 × 1010 CFU/day, in fermented juice (pineapple, banana, and papaya peel) GD90–PND28 |
↓ Number of deaths before weaning ↑ body weight from PND7 until PND28 PND28: ↓ Diarrhea incidence, ↑ serum concentration of Na + , pCO2, and D-β-hydroxybutyrate, leukocytes, lymphocytes, and platelets in the piglets |
Not assessed | [72] | |
Large White × Landrace sows ± probiotics; offspring ± probiotics (split-plot design) Dams n = 12/group Offspring n = 10–6/group (male and female) |
Bacillus altitudinis WIT588 4 × 109 CFU/day (gestation), 1.2 × 1010 CFU/day (lactation), 1 × 109 CFU/day (offspring) in feed GD100–PND26, PND26–PND54 (offspring) |
PND21: No effect on body weight PND105, PND126: ↑ body weight PND34: Duodenum: ↑ villus length and area, crypt depth; Jejunum: ↑ crypt depth and area |
Dams Strain detected in feces Offspring PND13: Strain detected in the feces of 12/20 piglets PND26: Strain detected in 16/20 piglets |
[68] | |
Landrace x Yorkshire sows ± low/high-dose probiotics Dams n = 5/group Offspring n = 55/group (male and female, n = 5/group for microbiota) |
Enterococcus faecium DSM 7134 2.7 × 107 CFU/kg (low-dose group), 5.4 × 108 CFU/kg of feed (high-dose) GD101–PND21 |
↓ piglet pre-weaning mortality PND7: ↓ Diarrhea score PND21: ↑ Weaning weight, average daily gain and gain:feed ratio |
Offspring PND21: ↑ Fecal Lactobacillus and Enterococci, ↓ Escherichia coli counts PND35: ↑ Fecal Lactobacillus and Enterococci |
[71] | |
Atopic diseases | |||||
BALB/c mice exposed to ROFA or PBS on GD14, 16, 18 (air pollution exposure) Neonatal asthma model (ovalbumin sensitization) ± probiotics Dams Undisclosed Offspring n = 6–21/group (male and female) |
Bifidobacterium breve M-16 1 × 109 CFU/day in feed GD14–PND21 |
PND30: ↓ Eosinophils in the bronchoalveolar lavage fluid; ↓ allergic lung inflammation; ↓ mucus production; ↓ IL-5, IL-13, and Muc5ac gene expression |
Offspring PND30: ↓ Firmicutes proportion ↑ Gemella, Streptococcus, Dehalobacterium & ↓ Lactobacillus |
[81] | |
BALB/c mice ± probiotics AD induced with MC903 or not (offspring, PND21) Dams n = 6/group Offspring n = 8/group (male and female) |
Limosilactobacillus reuteri Fn041 or Lacticaseibacillus rhamnosus GG 1 × 109 CFU/day by gavage GD14–PND21; PND21–PND28 (offspring) |
PND28: Both strains: ↓ AD symptoms (skin swelling, mast cell, eosinophil infiltration), ↓ IL-4 and IL-12; L. reuteri Fn041: ↓ IgE, ↑ proportion CD4 + CD25 + Foxp3 + Tregs cells in mesenteric lymph nodes |
Offspring PND28: L. reuteri Fn041: ↑ Actinobacteria, Lactobacillus and Akkermansia; LGG: ↓ ⍺-diversity; ↑ Escherichia_Shigella, Enterococcus; Both strains: ↓ Alloprevotella, Klebsiella, Helicobacter |
[82] | |
Inflammation | |||||
C57BL/6 mice ± probiotics, DSS induced inflammation at PND70 Dams n = 2/group Offspring n = 4/group (male, microbiota) n = 15–16/group (female, weight loss–sample size microbiota undisclosed) |
Mixes: Limosilactobacillus reuteri (strains 6798‐1, 6798‐jm, and 6798‐cm) or Lactobacillus johnsonii (strains 4901, 4903, 4931) 1 × 109 CFU/ml by gavage GD12–birth; PND7–PND21 (every other day) |
PND76: L. reuteri: ↓ post-DSS weight loss in females |
Offspring Females: Both strains, PND21: Altered Muribaculum and Ruminiclostridium abundance PND30: Significant effect on β-diversity, altered Lachnoclostridium, Muribaculum, and Tyzzerella abundances L. reuteri, PND70: ↓ Lachnoclostridium, Akkermansia, Parasutterella, Ruminiclostridium, Bacteroides, ↑ Lactobacillus, Muribaculum, and Bifidobacterium Males: L. reuteri, PND70: ↓ Alistipes abundance, no effect on female altered genus |
[73••] | |
SPF C57BL/6 J mice ± probiotics, IL-1β induced postnatal systemic inflammation (PND14 and 28) Dams n = 5–6/group Offspring n = 3–9/group/time point (male and female) |
Mix: Lactobacillus acidophilus ATCC 53544, Bifidobacterium infantis ATCC 15697 1 × 109 CFU/day each GD16–PND21 Undisclosed |
PND14: ↓ IL-1β-induced systemic levels of IL-6, KC, MCP-1, and IL-1β; normalize blood–brain-barrier permeability and occludin expression; regulate recruitment and extracellular matrix damage Promote neuronal development (↑ NeuN, NFL, and Syn1 expression) and oligodendrocyte progenitor cell development (↑ NG2 expression) PND28: No effect on IL-1β-induced ↑ of MCP-1 levels |
Not assessed | [113] | |
Obesity | |||||
CD-1 IGS mice receiving a HFD or not started 6 weeks before mating until PND21, ± probiotics Dams n = 4/group Offspring n = 10–15/group/time point (male) n = 9–14/group/time point (female) |
Mix: Bacillus subtilis PXN®21®, Bifidobacterium bifidum PXN® 23™, Bifidobacterium breve PXN® 25™, Bifidobacterium infantis PXN® 27™, Bifidobacterium longum PXN® 30™, Lactobacillus acidophilus PXN® 35™, Lactobacillus delbrueckii ssp. bulgaricus PXN® 39™, Lactococcus casei PXN® 37™, Lactiplantibacillus plantarum PXN® 47™, Lacticaseibacillus rhamnosus PXN® 54™, Lactobacillus helveticus PXN® 45™, Ligilactobacillus salivarius PXN® 57™, Lactococcus lactis ssp. lactis PXN® 63™, Streptococcus thermophilus PXN® 66™ Total of 4 × 107 CFU/ml in drinking water GD0.5–PND21 |
PND21: ↑ Female and male body weight; altered expression of gene involved in synaptic plasticity: restored HFD-induced ↑ of GLUN2C in female and modulate genes expression in control and HFD groups (GLUN2A, PFKB3, female GLUN2B, male SYP and CREB1) ↑ IL-6 prefrontal cortex expression in control and HFD, ↓ liver TLR4, and liver IL-1B in males PND21 & PND112: Prevent HFD-induced anxiety-like behavior ↑ gut propionate, butyrate, and brain lactate in control and HFD PND112: ↑ Brain-derived neurotrophic factor expression (BDNF, PFKB3, ΔFOSB and male cFOS and ZIF-268), ↑ female liver IL-6 in control and HFD |
Dams PND21: ↑ Fecal Lactobacillus spp. in control and HFD, and ↑ Bifidobacterium spp. in control diet group |
[79••] | |
Sprague Dawley rats receiving HF diet or not from GD1 until PND21, ± probiotics Dams n = 3/group Offspring n = 7–8/group (male) |
Lacticaseibacillus casei 2 × 108 CFU/day by gavage GD1–PND21 |
12 weeks: No effect on body weight, Prevents HF-induced hypertension: ↓ systolic blood pressure from 6 to 12 weeks. ↓ acetate plasma levels; ↓ renal mRNA expression of Olfr78 |
Offspring 12 weeks Compared to control: ↑ Parabacteroides and Akkermansia muciniphila, ↓ Actinobacteria, Bacteroides, Bacteroides/Prevotella, Actinobacteria/Firmicutes ratio Compared to HF group: ↑ Lactobacillus, Acholeplasma, Bacteroides acidifaciens, ↓ Alkaliphilus, Leptolyngbya, Prevotella albensis, Ruminococcus albus |
[95] | |
Sprague Dawley rats receiving HFD or not from GD1 until PND21 ± probiotics Dams n = 3/group Offspring n = 8/group (male) |
Lacticaseibacillus casei 2 × 108 CFU/day by gavage GD1–PND21 |
16 weeks: Compared to HFD: No effect on BW, ↓ systolic blood pressure from 12 to 16 weeks, ↓ fecal propionate level, ↓ plasma TMAO levels and TMAO-to-TMA, ↑ DMA-to-TMAO conversion ratio, ↓ renal mRNA Ace expression |
Offspring 3 weeks: Compared to HFD: No effect on ⍺- or β-diversity; Prevent the HF-diet induced ↓ Alkaliphilus, Akkermansia abundances and several Lactobacillus species 16 weeks: Prevents the ↓ Lactobacillus |
[96] | |
Wistar rats receiving HFHC diet or not from GD1 until PND21 ± probiotics Dams n = 5/group Offspring n = 5–10/group (male) |
Lactiplantibacillus plantarum WJL 1 × 109 CFU/day by gavage GD1–birth; PND2-21 |
↓ body weight and length from PND30 to PND90 ↓ blood pressure and recovered vascular function in later life |
Dams Restore ⍺-diversity but not β-diversity No effect on dyslipidemia induced dysbiosis ↑ Lachnospiraceae (Ruminococcus, Blautia, Dorea, and Coprococcus) |
[94] | |
Neurodegenerative disease | |||||
Wistar Albino rats receiving intraperitoneal injection of LPS (100 μg/kg) on GD17 (neuroinflammation) ± probiotics Dams n = 5/group Offspring n = 18/group (male and female) |
Ligilactobacillus salivarius ATCC 11741, or B. bifidum ATCC 29521, or both Total of 4 × 109 CFU/ml by gavage GD1–birth |
PND1: Improved brain APP levels and mRNA, gamma-secretase and beta-secretase levels, improved brain BDNF mRNA expression ↓ brain damage in the cortical area |
Not assessed | [114] | |
Sprague Dawley rats exposed to lead one week after mating until offspring sacrifice (memory dysfunction model) ± probiotics Dams n = 3/group Offspring n = 6–10/group (female) |
Mix: Bifidobacterium longum BL986, Lactobacillus acidophilus LA1063, Limosilactobacillus fermentum LF26, Lactobacillus helveticus LH43, Lacticaseibacillus paracasei LPC12, Lacticaseibacillus rhamnosus LRH10, Streptococcus thermophilus ST30 Total of 1 × 1010 CFU/day in drinking water GD7–PND21, PDN21–PND68 (offspring) |
PND22 & PND68: Reversed lead-led memory impairment and loss of hippocampal spines PND68: Restore IL-6 levels and the epigenetic event H3K27me3 level in the hippocampus |
Offspring PND68: Restore ⍺-diversity, Firmicutes/Bacteroidetes ratio, Proteobacteria, Actinobacteria. ↑ Helicobacter, Bifidobacterium, Bacteroides, ↓ Anaerovibrio, Ruminococcaceae_UCG-008, Lactobacillus |
[100] |
AD atopic dermatitis, BMI body mass index, BW birth weight, CAT catalase, CFU colony forming units, FOS fructo-oligosaccharides, GD gestational day, GPx glutathione peroxidase, HDL high-density lipoprotein, HF high-fructose, HFD high-fat diet, HFHC high-fat and high-cholesterol, Ig immunoglobulin, IL interleukin, LDL low-density lipoprotein, LPS lipopolysaccharides, MOS mannan-oligosaccharides, MUC2 mucin 2, PND postnatal day, ROFA residual oil fly ash, SCFA short-chain fatty acids, SI small intestine, SOD superoxide dismutase, SPF specific pathogen-free, ZO-1 zonula occludens 1