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Table 1 Summary of the animal studies investigating the effects of fasting on gut microbiota alterations

From: Gut microbiota modulation as a possible mediating mechanism for fasting-induced alleviation of metabolic complications: a systematic review

First author, year Animal model Intervention Biospecimen Microbiota analyzing methods Main findings
Sonoyama K, 2009 Male Syrian hamsters (age: 10 weeks) 1. Fed active group (n = 6) Cecum 16S rRNA clone library and species-specific real-time quantitative PCR ↑Akkermansia muciniphila, a mucin degrader, in fasting group but not in hibernation
   2. Fasted active, group (n = 6) fasted 96 h    ↑Clostridia in fed active and hibernating group
   3. hibernating group (n = 6) maintaining darkness at 4 °C    
Kohl KD, 2014 Fish, toads, geckos, quail, mice Four time points through Colon & cecum 16S rRNA gene sequencing In tilapia, quail, and mice: ↓Lactobacillus & Prevotella, ↑ Oscillospira
   28 days fasting    In tetrapods ↓ Coprobacillus & Ruminococcus
Li G, 2017 7–8-week-old male C57BL/6 N mice: diet-induced obese model 1. Controls: fed with ad libitum (AL) diet Cecum 16S rRNA gene amplicon sequencing ↑Firmicutes, ↓ Bacteriodetes and Actinobacteria, ↑ Firmicutes to Bacteroidetes ratio
   2. Intermittent fasting (IF): every other day fasting (EODF) regimen    ↑ beiging of white adipose tissue and subsequently ameliorate metabolic disorders
McCue MD, 2017 Mice, quail, tilapia, toad, geckos Mice (3 days fasting), quail (7 days fasting), tilapia & toad (21 days fasting), geckos (28daysfasting) Feces 16S rRNA sequencing Alteration in Bacteriodetes, Firmicutes, Proteobacteria, Fusobacteria and Verrucomicrobia
Beli E, 2018 4-month-old db/m (non- diabetic) and db/db (diabetic) mice 1. Controls: fed with ad libitum (AL) diet Feces 16S rRNA sequencing with the MiSeq platform ↑ Firmicutes, ↓ Bacteroidetes and Verrucomicrobia, ↑gut mucin, goblet cell number, villi length
   2. IF: fasted on an every other 24-h interval for 7 months    ↓ plasma peptidoglycan, ↑ tauroursodeoxycholate bile acid ↓retinal TNF-α mRNA
Catterson JH, 2018 fruit flies (Drosophila melanogaster) 1. Controls: fed with ad libitum Blood and tissue qPCR quantification of bacterial load ↓age-related pathologies,↑ gut barrier function & gut health due to ↓relative bacterial abundance
   2. IF: 2-day fed,5-day fasted for 40 days    
Cignarella F, 2018 7- week- old Female C57BL/6 J mice: multiple sclerosis (MS) animal model 1. Controls: fed with ad libitum (AL) diet (n = 10) Feces 16S rRNA gene sequencing Lactobacillaceae, Bacterioidaceae and Prevotellaceae families
   2. IF: fasted, fed every other day (n = 10)    ↑ketone formation and glutathione metabolism, ↑ anti-oxidative pathways, ↓IL-17 producing T cells & ↑the number of regulatory T cells ↓inflammation, demyelination and axonal damage
Wei S, 2018 6-week-old male C57BL/ksJ-db (db/db) mice: a genetic model of type 2 diabetes 1. Controls: standard chow, free access to food and water Feces 16S rRNA gene sequencing ↑ Bacteroidetes, ↓Firmicutes & Saccharbacteria. ↑Parabacteroides & Blautia, ↓Prevotellaceae, Alistipes & Ruminococcaceae
   2. FMD: fasting with the (30% of the daily calorie intake of control group) for 1 week, followed by ad libitum feeding for another week    ↓ fasting blood glucose, ↓hepatic steatosis, ↓loss of pancreatic islets and β cells
Kim JN, 2019 Three ruminally cannulated Holstein steers 1.Controls: the ruminal fluids 2 h after the morning Rumen Denaturing gradient gel electrophoresis and quantitative polymerase chain reaction Anaerovibrio lipolytica, Eubacterium ruminantium, Prevotella albensis, Prevotella ruminicola, and Ruminobacter amylophilus
   2.Fasting: the ruminal fluids feeding and 24 h after fasting    
Li T, 2019 Crucian Carp fish 1. Controls: fed to satiation twice per day Gut 16S rRNA gene amplicon sequencing In IF groups: ↑alpha diversity and ecosystem stability of gut microbiota
   2.IF: fasted for 5 days followed by 5 days of re-feeding   Bacteroides, Akkermansia, and Erysipelotrichaceae, ↑growth performances, immune function
   3. Long term fasting (LF): not fed anything during the whole experimental period    In LF group: ↑ Proteobacteria, Gammaproteobacteria, Vibrio, other Vibrionaceae
Miyamoto J, 2019 Wild-type and Gpr43 − / − mice 1.IF: alternating 24 h periods (15 cycles for 1 month) of free access to diet followed by 24 h fasting Cecum 16S rRNA amplicon sequencing IF group: ↑Bacteroidetes& Verrucomicrobia
   2. Eucaloric ketogenic diet: ketogenic diets for 6 weeks    ↑total plasma ketone body ↓total plasma & cecal short chain fatty acids (SCFA)
Park S, 2019 Male Sprague Dawley rats: Alzheimer’s disease (AD) model 1. Controls: normal diet Feces 16S rRNA amplicon sequencing IF group: ↓ Clostridales, ↑Lactobacillales
   2. Ketogenic diet    Ketogenic group: ↑the relative counts of Proteobacteria especially Enterobacteriales
   3. High carbohydrate diet    IF and high carbohydrate groups, but not ketogenic group:
   4.IF: normal diet with intermittent fasting    ↓ the hippocampal amyloidβ deposition,↑ memory function
Rangan P, 2019 8-weeks-old female C57BL/6 mice: inflammatory bowel diseases mouse model 1. Controls: standard diet Feces 16S rRNA gene sequencing FMD: ↑ Microbial strains associated with Tcell regulation and gut regeneration
   2. Fasting-mimicking diet (FMD): control diet plus 2 cycles of FMD    ↓ intestinal inflammation, ↑stem cell number,
   3. water-only fasting (WF): control diet plus 2 cycles of WF   Bifidobacteriaceae, ↓Lactobacillaceae
      WF: ↑regenerative, ↓ inflammatory markers without reversing pathology
Zhou ZH, 2019 6-week-old male C57BL/6 J mice Parkinson’s disease (PD) model 1. Controls: normal diet (12 kcal/day) feces 16S and 18S rRNA gene sequencing FMD: ↑ Firmicutes, Tenericutes, Opisthokonta, ↓Proteobacteria
   2. Fasting-mimicking diet (FMD) day 1: 50% of the standard daily calorie intake day 2–3: 10% of the standard daily calorie intake    ↑neuroprotective effect for PD
Deng Y, 2020 3-week-old male C57BL/6 J mice:diet-induced obese model 1.controls: fed with ad libitum (AL) diet (n = 15) feces 16S rDNA gene amplicon sequencing ↑community diversity in gut flora
   2. IF: 24-h fasting for 30 days (n = 15)    ↓ Firmicutes to Bacteroidetes ratio
      ↓fat accumulation, ↑ white fat conversion to beige
Li L, 2020 7-week-old male C57BL/6JLvri mice 1. Controls: fed with ad libitum (AL) diet (n = 15) Feces next-generation sequencing ↑level of Akkermansia & ↓level of Alistipe in 16 h fasting group
   2. IF: divided in 3 groups of fasting: 12, 16 and 20 h (n = 15 per group)   of 16S r RNA gene amplicons ↓ cumulative food intake in the 16 and 20 h fasting groups
Liu Z, 2020 homozygous Leprdb/db mice (diabetic), heterozygous Leprdb/m, mice (non- diabetic) 1.controls: fed with ad libitum (AL) diet Feces 16S rRNA gene v3–v4 amplicon ↑ villi length and the muscularis thickness. ↓ gut leakage, ↑cognitive function
   2. IF: deprived of food for 24 h for 28-day   Sequencing ↑ microbiome alpha diversity
     Lactobacillus and butyrate-producing Odoribacter. ↓Enterococcus, Streptococcus, Candidatus Arthromitus, Rummeliibacillus, Enterococcaceae, Leuconostocaceae
Ye Y, 2020 8-week-old male Kunming mice 1. High-fat diet ad libitum Feces 16S rRNA gene V3–V4 ↓ weight gain, ↓liver steatosis, ↓hepatic levels of triglycerides
   2. Time-restricted high-fat diet restricted to an 8-h temporal window per day for 8 weeks   Sequencing ↑ Bacteroidetes, ↓ Firmicutes
Zhang X, 2020 7–8 week –old male C57BL/6 mice: colitis mouse model 1.Controls: standard diet Feces 16S rRNA gene ADF, TRF & IER: ↑Escherichia
   2. Alternate-day fasting (ADF): 24 h feeding/fasting   sequencing TRF and IER: ↓Escherichia
   3. Time restricted feeding (TRF): fed 8 h per day, fed at 24:00 p.m. and fasting at 8:00 a.m. in the morning    TRF and IER, but not ADF: ↓Gammaproteobacteria, Enterobacteriaceae, Shigella & Escherichia coli,Christensenellaceae
   4. Intermittent energy fasting (IER): two cycles of four days of IER diet from day 11–14 and day 29–32    ↑SCFAs generation-related microbes: Rikenellaceae, Lactobacillus, Coproccus & Ruminococcus
      IER but not TRF and ADF: ↓Peptostreptococcaceae
Shi H, 2021 5 weeks old, WKY and SHRSP hypertensive stroke-prone rats 1. Controls: ad libitum feeding 2.every other day fasting (EODF) group: alternating 24 h of ad libitum food access followed by 24 h fasting for 10 weeks Cecum & plasma Shotgun sequence analysis of the microbiota and untargeted metabolomics ↑Bacteriodetes & Actinobacteria, ↓proteobacteria
      ↑ microbial bile acid metabolism genes: 7α-dehydroxylase and bile salt hydrolase
      ↓ body weight & systolic blood pressure