We retrieved a total of 2864 references. After eliminating duplicates (996) with the bibliographic tool Mendeley it remained 1868 references. 73 studies fulfilled the criteria for inclusion. The search strategy and selection of publications are shown in Figure 1.
[Here Figure 1]
Of the total of 73 studies, 50 are research studies, and 22 are reviews or meta-analysis papers. Every research study found was classified into one of the following four categories of factors:
-DIET: 22 papers addressing probiotics, prebiotics, sweeteners, type of foods, ultra-processed food, specific diets (Mediterranean diet, vegan diet), fermented foods, plant-based food polyphenols, or protein supplements (Table 1)
-HABITS: 7 studies about sleep, physical activity, tobacco, alcohol, or drugs (Table 2).
-ENVIRONMENT: 11 articles regarding rural/urban environment, living near a mine, nursing homes, cohabiting family (partner, children, parents, grandparents), short hospitalization periods, long ocean voyages, or exposure to high altitudes (Table 3).
-CHEMICALS: 10 papers about antibiotics, methamphetamine, opioids, metformin, vitamin D, proton-pump inhibitors, herbicides, fungicides, titanium oxide nanoparticles, or herbal treatments (, Table 4).
One study addressed two or more of these categories and was included in the discussion.70
First, a review was made of the results related to the factors studied in every paper. In 41 of these research studies questionnaires were used. In Tables 1 to 4, questions of the questionnaires used in every study are split into the four categories mentioned (diet, habits, environment, and chemicals) and data of these four categories retrieved in every study were recorded.
Diet
Overall, diet was the most studied factor about its influence on digestive microbiota (43% of the papers included in the scoping review) (Table 1). In the literature reviewed, probiotics have been shown to influence digestive microbiota to prevent or treat some diseases, decrease dysbiosis due to age, or help to maintain a normal microbiota in a meat-based diet. 13, 14. 15 Besides, the effect of commercial probiotics in healthy subjects was proven to increase the diversity of the oral cavity.16 Fermented food has taken greater importance due to its relationship with positive microbiota changes and other effects such as reduction of academic stress and inflammatory markers. 17, 18, 19
The intake of dietary fibre with different prebiotics also influences digestive microbiota. A diet rich in fiber increases the levels of beneficial bacteria and short-chain fatty acids (SCFA). 20, 21, 22 Supplementing with milk oligosaccharides, a prebiotic based on resistant starch, induces the growth of beneficial bifidobacteria. 23, 24 It has also been described that a greater intake of dietary fiber from cereals, fruits, and vegetables is associated with shifts in gut microbiome composition and a reduction of C-reactive protein. 25 Inulin consumption modified the microbiota composition with higher microbial diversity 26. Dietary addition of L. casei, B. breve and a prebiotic product, results in an improvement of intestinal physiologic traits and an increase of total fecal bifidobacteria 27.
Polyphenols also have been shown to affect the gut microbiota. Some of them have protective effects of the goblet cells of the intestinal mucosa. Other polyphenols have been shown to increase the counts of Lactobacillus and Bifidobacterium and improve gut and host health. 28, 29 The type of diet has also been related to gut microbiota status. High adherence to Southern European Atlantic Diet produces a greater number of total bacteria, Actinobacteria, Lactobacillus, and Bifidobacterium genus.30 Significant correlations between bacterial taxa and Mediterranean Diet adherence were observed in centenarians, nonagenarians, and younger subjects.31 Microbiota associated to Mediterranean diet was enriched with beneficial bacteria that induces anti-inflammatory status and reduces the risk of developing gut diseases. 32 Individual foods have also been shown to influence digestive microbiota. Green tea liquid consumed daily decreased Bacteroidetes in feces and pathogenic bacteria in saliva. 33 A diet heavily based on beans and dairy products reduced gut diversity with less beneficial bacteria.34 A long-term protein supplementation may have a negative effect on gut microbiota and increase the risk of intestinal diseases.35, 36 Nevertheless, natural plant extracts with polyphenols, prebiotics, probiotics, and regular physical exercise improved gut microbiota, reducing dysbiosis produced by High Protein Diet (HPD). 36
Consumption of more than five servings per day of ultra-processed food produced a decrease in microbiota diversity 37. Sesame seed snacks and plant-based smoothies produced a notable increase in endogenous antioxidants in plasma and an improvement in gut microbiota. 38
Pure saccharin supplementation did not alter gut microbial diversity, while Vitamin D supplementation increased the overall diversity of gut microbiota in healthy females with vitamin D deficiency. 39, 40 Food additives also affect gut microbiota composition. Some sweeteners and some emulsifiers may alter microbiota. Food preservatives increase the growth of proinflammatory bacteria, while anti-inflammatory bacteria decrease. Acidifiers can reduce coliform presence and increase lactic acid bacteria, and colorants can reduce microbiota diversity in vitro. Nevertheless, the long-term impact of food additives on gut microbiota needs to be examined 41.
[Here Table 1]
Habits
As shown in Table 2, there are different types of habits that influence digestive microbiota, such as physical activity, which has been shown to increase microbial diversity. 42 The quality of sleep also influences the microbiota, although not in all the circumstances. The microbiome is resistant to changes after consecutive days of sleeping restriction. 43 However, an acute sleep-wake cycle shift of 2 to 4 hours (subjects postpone their regular sleeping time for 2 to 4 hours) affects the functional profiles of gut microbes and interactions among them 44. Alcohol consumption or smoking or the combination of both cause gut microbiota dysbiosis in healthy men. Noteworthy, smoking alone causes more severe dysbiosis than drinking alone. 45 Furthermore, smoking habits also modified oral microbiota diversity and heavy episodic drinking is associated with a specific stool type phenotype (Bristol stool scale). 46 Consumption of heroin, ephedrine, and methamphetamine produced changes in the microbiota. Bacterial diversity was higher in SUD (substance use disorders) and this increases with the length of substance abuse.47 Intestinal colonization by oral bacteria has been demonstrated; therefore, oral hygiene becomes more important 46,48. In fact, the salivary microbiome was affected by denture use 46.
[Here Table 2]
Environment
The environment is also an important factor to study in this context (Table 3). References retrieved have shown different sources of influence on gut microbiota. Blastocystis, a component of gut microbiota, has a higher prevalence in non-westernized individuals 49. Differences in gut microbiota composition were found in individuals with the same ethnicity but coming from different Italian regions. 50 Medication and time spent in the nursing home contributed to microbiota dysbiosis more than age and frailty. The microbiota of residents who had lived in the nursing home for more than 1 year were enriched in inflammatory and pathogenic species and reduced in anti-inflammatory and symbiotic species. 51
Nevertheless, it has been described that short-term hospitalization did not impact the richness or structure of the salivary microbiome.52
Indoor spaces often harbor unique microbial communities. Indoor air in residences contained a greater number of culturable bacteria than outdoor air. Differences in bacterial flora in settled dust in two buildings were greater than differences between seasons. Bacterial communities on surfaces in offices were different between cities, and the microbiota in a home were identifiable by family. Humans are also major sources of bacteria in indoor air. 53
As for the quality of the environment, long-term exposure to metals in residents near a mining and smelting area produced different profiles of gut microbiota. These effects were higher in men because of mining and smelting activity.54 Currently, it is under discussion whether air pollution due to emissions from factories, chimneys, or livestock also affects gut microbiota since it may be trapped in the food. Some studies showed that gut microbiota metabolized the inorganic arsenic into toxic metabolites. Currently, there are some chemicals and pesticides that are considered microbiota-disrupting chemicals. 55 The living altitude also influences the microbiota composition, which was enriched in Prevotella and Bacteroidetes in subjects living in high altitudes (>1500m) while Faecalibacterium and Blautia were increased in the low altitude group (<1500m). 56
Microbiota diversity can also change due to traveling. Sun et al., (2022) reported that a 135-day sea expedition produced “seafaring syndrome” (SS) with abnormal defecation, frequent insomnia, poor sleeping quality, and overeating. Noteworthy, a significant correlation between the gut microbiome perturbation and this syndrome was found. 57
Cohabitation with other people also influences digestive microbiota. Microbial dysbiosis might be caused in healthy partners cohabiting with Ulcerous Colitis patients and anxiety and stress are related to reduced diversity. 58, 59 People with larger social networks tend to have a more diverse microbiome. Cohabiting time affected strain sharing more than age or genetics. 60 A high similarity in the oral microbiota composition was found between members of the same family, higher between twins, parents, or brothers, and less or none between grandparents and grandchildren. There was also a similarity between classmates. 9
[Here Table 3]
Chemicals
Chemicals have proven to be important factors that influence digestive microbiota (Table 4). Non steroidal anti-inflammatory drugs (NSAID) provoke changes in gut microbiota. Bacterial population in the gastrointestinal tract reflects the combination of medications and varied with the type of NSAID that people take 61. Proton Pump Inhibitors (PPI) have been associated with a decrease in microbiota diversity and an increase of Lactobacilus, Steptococcus, Staphylococcus and E.coli. 62, 63 Antibiotics are clearly an important factor that changes gut microbiota characteristics. Azithromycin alters the diversity of the salivary microbiome and delays the recovery of the diversity of gut microbiota after a treatment. 64 Moxifloxacin reduced bacterial diversity, which is not recovered until 16 to 21 days 65. S. boulardii treatment can mitigate some antibiotic-induced microbiota changes (dysbiosis).66 Metformine has an effect of reduction of gut microbiome diversity already after 24 hours of administration.67 Decreased diversity of Shanon and Chao indices was observed in methadone-treated subjects. 68 Also, essential oils may be considered as a factor that influences the gut microbiome. Notably, a component of these oils, geraniol, has proven to increase butyrate-producer bacteria and improve intestinal dysbiosis.69
[Here Table 4]
Diet, chemicals, environment, and/or habits interaction
Summing up, diet, chemicals, environment, and habits have been described as influencing the digestive microbiota. These factors are typically considered individually without considering the interactions between them. Diet, sleep quality, medications, supplements, physical activity, and psychological status are considered possible causes of dysbiosis. 70, 71, 72 However, it is also very important to study the interactions between factors on the microbiota quality. Boisseau et al. (2022) found that the microbiota of athletes is different from the rest of the population. Diets and treatments could be the origin of these differences. However, it is also known that exercise improves microbiota in sedentary people, but excessive practice does not improve microbiota and even worsens health. Nevertheless, there are no studies on humans that search for the interaction between diet and physical activity. 6 Also, a positive interaction was found between diet, physical exercise, and HPD. Natural plant extracts with polyphenols, prebiotics, probiotics, and regular physical exercise improved gut microbiota, reducing dysbiosis produced by HPD 36.