Streptococcus thermophilus

Streptococcus thermophilus: A Comprehensive Reference

1. Identity, Taxonomy, and Microbiology

1.1 Scientific Name and Taxonomy

Streptococcus thermophilus (abbreviated S. thermophilus or STH) is a Gram-positive, lactic acid–producing bacterium. It is a non-pathogenic, homofermentative facultative anaerobe with a long history of use in the home-made and modern industrial manufacture of fermented dairy products, especially yogurt. Taxonomically, the species belongs to the family Streptococcaceae and to the broader salivarius group of streptococci. During the period 1984–1995, S. thermophilus was classified as a subspecies of Streptococcus salivarius (S. salivarius subsp. thermophilus), but later studies based on 16S rRNA gene comparisons showed that, although closely related to S. salivarius, S. thermophilus is a distinct species.

Morphologically, S. thermophilus is cytochrome-, oxidase-, and catalase-negative, nonmotile, nonspore-forming, and homofermentative, existing in the form of cocci in pairs and chains. It is classified as a thermophile, growing optimally at 45°C with no growth at 10°C. The genus name "Streptococcus" derives from a Greek term meaning "twisted kernel," referring to the way the bacterium is grouped in chains that resemble a string of beads.

1.2 Genomic Diversity and Evolutionary Origin

STH is a clonal species that emerged only recently on the evolutionary timescale (3,000–30,000 years ago), from a commensal ancestor of the salivarius group. Its remarkable adaptation to grow in milk — a narrow and well-defined niche — has resulted in genome shaping through loss-of-function events and horizontal gene transfers (HGT). At least 26 strains of S. thermophilus have been identified and had their genomes sequenced. The low occurrence in S. thermophilus genomes of genes coding for biogenic amine production and antibiotic resistance is a contributing factor to its safety status.

1.3 Natural Sources and Ecological Niche

Streptococcus thermophilus is a natural inhabitant of raw milk in many parts of the world. Beyond raw milk, STH is also found in dairy products spontaneously fermented, such as diverse fermented milks in Mongolia and Ragusano cheese in Sicily. In addition, it can participate in the fermentation of leguminous products such as soy juice and milk kefir grains.

1.4 Common Forms and Preparations

S. thermophilus is encountered in several distinct forms:

• Live-culture fermented dairy foods: Streptococcus thermophilus is a beneficial lactic acid–producing bacterium widely used in the production of fermented dairy products such as yogurt, kefir, and cheese.
• Probiotic dietary supplements: S. thermophilus is commonly used to produce fermented dairy foods such as yogurt and is also used in probiotic supplements. Many commercial probiotic products available on the market all over the world contain STH.
• Topical (skin-care) preparations: Experimental creams containing sonicated Streptococcus thermophilus have been used to increase skin ceramide levels in healthy subjects.
• Infant formula supplements: S. thermophilus has been incorporated into infant formula products, sometimes in combination with other probiotic strains.

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2. Historical and Traditional Use

2.1 Pre-Modern Fermentation Traditions

S. thermophilus has been used for centuries — indirectly — through the fermentation of dairy products in Mediterranean, Central Asian, and Middle Eastern cultures. Traditional yogurt and kefir-making practices naturally cultivated this organism without awareness of its precise microbiological identity. Modern microbiology began isolating and characterizing S. thermophilus in the early 20th century, particularly as food science advanced and commercial fermentation processes were standardized.

S. thermophilus has been used to make yogurt and similar fermented dairy foods for centuries, including skyr, a staple of Icelandic diets since the Viking age. Like other probiotics with an affinity for dairy, S. thermophilus' purpose in these traditions was to turn lactose — the sugar in milk — into lactic acid, transforming milk into the gel-like, metabolite-rich substance humans have enjoyed since ancient times.

2.2 Industrial Era and Scientific Characterization

S. thermophilus is one of the most widely used bacteria in the dairy industry. USDA statistics from 1998 showed that more than 1.02 billion kilograms of mozzarella cheese and 621 million kilograms of yogurt were produced from S. thermophilus. The isolation and scientific characterization of the organism accompanied the industrialization of yogurt manufacturing in the 20th century, when the stable co-culture of S. thermophilus with Lactobacillus delbrueckii subsp. bulgaricus was formalized as the standard yogurt starter.

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3. Key Constituents, Metabolites, and Active Compounds

3.1 Lactic Acid and Fermentation Products

As a dairy starter, S. thermophilus can rapidly convert lactose into lactic acid, which causes a rapid reduction in pH resulting in coagulation of milk proteins (casein). It ferments lactose homofermentatively to give L(+) lactic acid as the principal product. Aromatic compounds such as acetaldehyde and phenylacetic acid are products of amino acid catabolism and carbohydrate metabolism, playing a significant role in the sensory characteristics observed in dairy fermentations.

3.2 Beta-Galactosidase (Lactase)

One of the most important properties of this bacterium is the production of lactase, an enzyme that converts lactose (milk sugar) into simple sugars, which helps people who are lactose intolerant to digest milk. This enzyme is central to the organism's probiotic value for digestive health.

3.3 Exopolysaccharides (EPS)

Certain strains of S. thermophilus are capable of producing exopolysaccharides (EPSs) — high-molecular-weight polymers composed of repeating sugar units, synthesized and secreted by the bacterium. S. thermophilus exopolysaccharides have received wide attention over recent decades because they can improve the properties of the dairy product and confer beneficial health effects. EPS production is strain dependent, with the majority of strains producing between 20 and 100 mg of polymer dry mass per liter of fermented milk medium. EPS display strain-specific molecular weights ranging from 10 to 2,000 kDa and contribute to the viscosity of fermented products, while also providing antioxidant and immunomodulatory benefits. EPSs from dairy starters not only show well-documented texturizing functionalities but are also fermentable substrates, termed "prebiotics," for modulating the human gut microbiome.

3.4 Bacteriocins (Thermophilins)

Bacteriocins produced by S. thermophilus, such as thermophilins (e.g., Thermophilin 13, 110), exhibit extensive antimicrobial efficacy against pathogens including Listeria monocytogenes and Bacillus cereus. S. thermophilus is able to synthesize thermophilins, which are small peptides able to inhibit the growth or kill closely related bacteria; ten thermophilins have been identified from ten different strains of S. thermophilus. Their activity is modulated by quorum-sensing mechanisms involving the blp gene cluster, and they possess significant stability under heat and pH variations, making them suitable for biopreservation applications.

3.5 Folate (Vitamin B9)

Many S. thermophilus strains are able to produce and release folate during growth. Among sixteen commercial probiotic strains, S. thermophilus IDCC 2201 was identified as a major folate producer in metabolite profiling studies. Folate (vitamin B9) is involved, as a cofactor, in many essential functions of the cell such as synthesis of nucleic acids and amino acids, cellular growth, and cell division.

3.6 Gamma-Aminobutyric Acid (GABA)

Gamma-aminobutyric acid (GABA) is the main inhibitory neurotransmitter naturally occurring in the central nervous system and some peripheral tissues, with important physiological functions including neurotransmission, diuretic effects, and relaxation. Certain S. thermophilus strains are GABA-producing, capable of fermenting glutamate to produce GABA, which is also beneficial for improving the acid tolerance of the strain.

3.7 Sphingomyelinase

The presence of high levels of neutral sphingomyelinase activity in S. thermophilus is responsible for an observed increase of stratum corneum ceramide levels, leading to an improvement in skin barrier function and maintenance of stratum corneum flexibility. This enzyme is relevant to the organism's topical dermatological applications.

3.8 Urease

Among lactic acid bacteria of dairy interest, S. thermophilus is the only species harboring an active urease. The role of urease activity in modulating environmental pH and its involvement in the biosynthesis of aspartate, glutamine, and arginine have been investigated. Hydrolysis of urea increases the catabolic yield of S. thermophilus by modulating intracellular pH and increasing the activity of β-galactosidase, glycolytic enzymes, and lactate dehydrogenase.

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4. Mechanisms of Action

4.1 Lactose Digestion

S. thermophilus improves lactose tolerance primarily through two mechanisms: it reduces the lactose content of fermented dairy foods during fermentation, and it delivers active beta-galactosidase into the small intestine, where the enzyme can continue to hydrolyze residual lactose. Because S. thermophilus actively breaks down lactose during fermentation, live-culture yogurts and fermented milks typically contain less lactose than the original milk and deliver additional beta-galactosidase into the small intestine. Clinical work shows that many lactose-intolerant individuals tolerate yogurt better than equivalent amounts of unfermented milk, with fewer symptoms like bloating, cramps, and diarrhea.

4.2 Gut Barrier and Microbiome Modulation

Research has shown that S. thermophilus adapts its metabolism to stressful conditions found in the gastric and colonic competitive environment, and modifies its surface proteins during adhesion to intestinal epithelial cells. One plausible mechanism of pathogen exclusion is the co-aggregation of probiotics with pathogenic bacteria, which could prevent the attachment of pathogens to the intestinal surface and impede their colonization.

4.3 Immunomodulation

In vitro research published in PLOS ONE examined the effects of S. thermophilus on gene expression in human peripheral blood mononuclear cells. The study concluded that S. thermophilus 285 is able to lower inflammation (C3), and the strain was proposed as a viable candidate for further pre-clinical and clinical studies for the management of inflammatory diseases.

4.4 Anti-inflammatory Activity via Endocannabinoid System

Preliminary in vitro findings suggest that S. thermophilus may exert both potential analgesic and anti-inflammatory effects by modulating the endocannabinoid system (ECS) and reducing the degradation of endocannabinoids, known to play a key role in immune regulation and inflammation. Evidence shows significant inhibitory activity of S. thermophilus on fatty acid amide hydrolase (FAAH) and monoacylglycerol lipase (MAGL) activity, suggesting an ability to influence skin conditions by modulating ECS and preventing endocannabinoid degradation. These findings are described by the researchers as preliminary and require in vivo confirmation.

4.5 Folate Cross-Feeding in the Gut

When S. thermophilus was co-cultured with individual species comprising the human gut microbial community, specific bacteria such as Bacteroides thetaiotaomicron, Veillonella parvula, and Ruminococcus faecis grew dependently on both folate and S. thermophilus, exhibiting 2.8-, 3.6-, and 3.9-fold increases in growth in the presence of folate.

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5. Scientific Evidence by Area of Use

5.1 Lactose Intolerance

Evidence strength: Moderate (strongest area of evidence for this organism)

Clinical studies have shown that S. thermophilus, particularly when used in combination with other probiotics like Lactobacillus delbrueckii subsp. bulgaricus, can help alleviate symptoms of lactose intolerance by aiding lactose digestion in the gut. Yogurt bacteria have been shown to have a favorable impact on digestive health, improving lactose digestion in lactose-intolerant individuals. The mechanism — delivery of active beta-galactosidase — is well characterized, and the lactose intolerance application represents the most scientifically supported use of S. thermophilus in humans. However, the majority of this evidence comes from studies using yogurt (a combination of S. thermophilus and L. bulgaricus), making it difficult to isolate the independent contribution of S. thermophilus alone.

5.2 Antibiotic-Associated Diarrhea (AAD)

Evidence strength: Preliminary to Moderate (confounded by multi-strain formulations)

In a study examining the efficiency of a probiotic drink containing S. thermophilus along with Lactobacillus casei and L. bulgaricus for the prevention of diarrhea associated with antibiotic use and that caused by Clostridium difficile, the formulation was found to reduce the incidence of antibiotic-associated diarrhea (AAD), with potential to decrease morbidity, healthcare costs, and mortality if used routinely in patients older than 50. Some research also suggests that S. thermophilus may help support the balance of gut microbiota, improve intestinal barrier function, and reduce the duration of antibiotic-associated diarrhea in children and adults. In virtually all of these studies, S. thermophilus was used as part of a multi-strain product, making strain-specific attribution difficult.

5.3 Rotavirus-Associated Diarrhea in Infants

Evidence strength: Preliminary (small trials, multi-strain formulations)

Rotavirus-induced diarrhea poses a worldwide medical problem causing substantial morbidity and mortality among children in developing countries. In a double-blind, placebo-controlled trial, infants aged 5–24 months who were admitted to a chronic medical care hospital were randomized to receive a standard infant formula or the same formula supplemented with a probiotic combination including S. thermophilus. Other work with fermented infant formulas using S. thermophilus together with Bifidobacterium breve suggests reductions in the severity or duration of acute diarrhea episodes and favorable changes in stool patterns and some immune markers. These findings require replication in larger, adequately powered trials.

5.4 Necrotizing Enterocolitis (NEC) in Premature Infants

Evidence strength: Preliminary (as part of multi-strain combinations; not proven for S. thermophilus alone)

A 2013 study reported that the combination of B. infantis, Streptococcus thermophilus, and Bifidobacterium lactis was effective in the reduction of NEC. Network meta-analysis suggests the preventive efficacy of Bifidobacterium and Lactobacillus spp., but even more for mixtures of Bifidobacterium, Streptococcus, and Bifidobacterium and Streptococcus spp. The Probiotics in Extreme Prematurity in Scandinavia (PEPS) trial, a large ongoing multicenter, double-blinded, placebo-controlled RCT, is examining the combination of Bifidobacterium infantis, Bifidobacterium lactis, and Streptococcus thermophilus in extremely preterm infants. However, studies comparing different strains head-to-head are lacking, and differences in inclusion criteria, dosage, strains, and primary outcomes in most trials are major obstacles to providing evidence-based conclusions.

5.5 Skin Health — Ceramide Levels and Atopic Dermatitis

Evidence strength: Preliminary (small uncontrolled trials, topical application)

A significant increase in skin ceramide levels was observed in 17 healthy subjects after treatment with a cream containing S. thermophilus. Topical treatment with an S. thermophilus-containing cream also increased ceramide levels and hydration in the skin of 20 healthy elderly women. A 2-week application of the cream, containing a sonicated preparation of S. thermophilus, in the forearm skin of 11 atopic dermatitis patients led to a significant and relevant increase of skin ceramide amounts, which could have resulted from the sphingomyelin hydrolysis through the bacterial sphingomyelinase. In all patients the topical application also resulted in the improvement of the signs and symptoms characteristic of AD skin, including erythema, scaling, and pruritus. These trials lacked control groups and had small samples, so results should be interpreted with caution.

5.6 Immune System Modulation

Evidence strength: Preliminary (mostly in vitro and animal studies)

Reviews of lactic acid bacteria, including S. thermophilus, describe several potential gut benefits including supporting mucosal barrier integrity and tight junction function, producing metabolites that can nourish beneficial commensals or inhibit pathogens, and helping restore microbiota balance after antibiotic-associated disruption. Summarized data suggest S. thermophilus may contribute to reduced gastrointestinal inflammation and improved host defense, including interactions with Helicobacter pylori and modulation of conditions like irritable bowel syndrome, although much of this comes from preclinical or combined-strain studies.

5.7 Colorectal Cancer — Exploratory Research

Evidence strength: Very preliminary (in vitro and animal studies only)

β-galactosidase secreted by S. thermophilus has been found to inhibit tumor progression by modulating oxidative phosphorylation and Hippo pathway kinases in cell studies. Additionally, S. thermophilus strains isolated from dairy environments have demonstrated anticancer activity, possibly linked to folate production. In mouse fibrosarcoma and neuroblastoma cells, S. thermophilus exhibited antitumor effects through the activation of T-lymphocytes; however, the precise mechanisms underlying these effects remain poorly understood. No human clinical trials on S. thermophilus and cancer have been published; all findings to date are from in vitro or animal models and should not be interpreted as clinical evidence.

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6. Gastrointestinal Survival and Probiotic Status

A key scientific debate surrounds whether S. thermophilus qualifies as a "probiotic" in the strict FAO/WHO sense, which requires demonstrated survival through the gastrointestinal tract. Even though the market offers many probiotic products containing S. thermophilus biomasses, the applicability of the term "probiotic" for this species is still questioned due to its sensitivity to gastrointestinal tract (GIT) transit conditions.

From the human clinical trials reviewed in a major scientific review of the bacteria's survivability, three of them found no viable S. thermophilus detected in fecal samples, while five identified some bacteria surviving the gastrointestinal tract. However, the scope of human clinical trials was still limited — most study participants ingested the bacteria from yogurt or a probiotic mix, and mostly fecal samples were used, meaning there is not yet enough data to understand if the bacteria survive in other parts of the digestive system.

Variability in GI survival capacity between S. thermophilus strains suggests that some strains have developed specific resistance mechanisms to the main stresses encountered in the digestive environment, such as acid pH and bile. Even if most S. thermophilus strains appear to be sensitive to acid pH and bile salts, human studies have established their ability to survive passage through the GI tract and transiently colonize while ingested.

S. thermophilus is considered a transient bacterium in the human intestine. Although it does not set up permanent residence, it exerts some positive influences on gastrointestinal health.

A methodological concern in this area has also been raised: in most intervention studies reviewed, the identification of S. thermophilus strains from stools was not carried out with the necessary taxonomic accuracy to avoid their misidentification with Streptococcus salivarius, a common human commensal and a species phylogenetically close to S. thermophilus.

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7. Body Systems and Health Areas

• Gastrointestinal system: Yogurt bacteria have been shown to have a favorable impact on digestive health, improving lactose digestion in lactose-intolerant individuals, increasing intestinal regularity and digestion, preventing diarrhea, and stimulating the gut immune system.
• Immune system: Health benefits associated with probiotics including S. thermophilus involve the prevention and treatment of diarrhea symptoms, the prevention of irritable bowel diseases, colitis, and necrotizing enterocolitis, as well as potential treatment of extra-intestinal pathologies such as atopic dermatitis, hypercholesterolemia, and allergies.
• Skin and dermatological system: Topical preparations have shown effects on skin ceramide production and atopic dermatitis symptoms (see Section 5.5).
• Neonatal and infant health: Used in combination products studied for NEC and infant diarrhea prevention.
• Colorectal oncology (exploratory): Only in vitro and animal evidence to date.

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8. Dosage Forms and Dosages Reported in Studies

The probiotic market contains many products marketed as probiotics that contain Streptococcus thermophilus strains at several dosages. Commercial probiotic products with various organisms range in CFU/dose from 2 to 450 billion. Specific dosages reported in the clinical literature include:

• Topical dermatology: A 2-week topical application of cream containing a sonicated preparation of S. thermophilus applied to the forearm skin of 11 patients led to significant increases in skin ceramide amounts. The precise bacterial concentration in these topical preparations is not standardized in current published literature.
• Infant NEC prevention: In the PEPS trial, enrolled infants receive Bifidobacterium infantis, Bifidobacterium lactis, and Streptococcus thermophilus (ProPrems®) diluted in 3 mL breastmilk or placebo (0.5 g maltodextrin powder) diluted in 3 mL breastmilk per day until gestational week 34.
• Dietary supplement forms: S. thermophilus appears in probiotic capsules, powders, and fermented food products, often as part of multi-strain preparations. No standardized monograph dosage exists for S. thermophilus in isolation as a dietary supplement.

Overall, health effects of probiotic strains are highly strain-dependent, and there is not a single probiotic strain that can confer all the benefits previously reported.

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9. Safety Considerations

9.1 Regulatory Safety Status

S. thermophilus is classified by the FDA as GRAS — Generally Recognized As Safe — a designation based on decades of safe human consumption and scientific evaluation. In the European Union, the European Food Safety Authority (EFSA) has granted it QPS (Qualified Presumption of Safety) status, a designation reserved for microorganisms with a well-established history of safe use and no known safety concerns at the species level. In the most recent EFSA QPS review cycle (to March 2025), a search of 122 scientific articles potentially relevant for the QPS evaluation of S. thermophilus was conducted; following analysis of titles and abstracts, no articles passed to the full article evaluation phase, and consequently the QPS status of S. thermophilus was not changed.

S. thermophilus has a very long history of use in the dairy industry without any identified health problem, and virulence-related genes are absent from its genome.

9.2 Safety in Immunocompromised Populations

Although probiotics in general are considered safe, there are concerns about their use in certain cases. Some people, such as those with compromised immune systems, short bowel syndrome, central venous catheters, heart valve disease, and premature infants, may be at higher risk for adverse events.

9.3 Antibiotic Resistance Genes

Although S. thermophilus holds GRAS (FDA) and QPS (EFSA) status for food applications, the presence of antibiotic resistance in some strains warrants evaluation of potential gene transfer to other bacteria, including pathogens. The low occurrence in S. thermophilus genomes of genes coding for antibiotic resistance is a contributing factor to its safety status, though this varies by strain.

9.4 Relationship to Pathogenic Streptococcal Species

Although its genus, Streptococcus, includes some pathogenic species, food industries consider S. thermophilus a safer bacterium than many other Streptococcus species; yogurt and cheese that contain live cultures of S. thermophilus are thought to be beneficial to health.

9.5 GI Tract Safety Surveillance

Large surveillance datasets do not signal S. thermophilus as a frequent culprit for adverse events, and clinical trials have not highlighted specific invasive risks with well-characterized strains.

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10. Evidence Limitations and Research Gaps

Several important limitations must be acknowledged when evaluating the body of research on S. thermophilus:

• In contrast with other lactic acid bacteria, the probiotic status of S. thermophilus remains still questioned.
• The scientific evidence supporting the probiotic status of S. thermophilus strains is often contradictory.
• The majority of human clinical trials use S. thermophilus in combination with other probiotic strains, making it impossible to attribute observed effects solely to this organism.
• Health effects are highly strain-dependent, and there is not a single probiotic strain that can confer all benefits previously reported.
• The natural intra-species diversity of S. thermophilus represents an interesting source for innovation, but also means that findings from one strain may not extrapolate to others.
• There have been enough studies to suggest that S. thermophilus may alleviate lactose intolerance, though most of the clear evidence came from animal studies.

References

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• Tarrah A, et al. "In vitro Probiotic Potential and Anti-cancer Activity of Newly Isolated Folate-Producing Streptococcus thermophilus Strains." Frontiers in Microbiology, 2018.
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