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Bacilli

Bacilli is a class of within the phylum (formerly known as Firmicutes), characterized by low G+C content (typically less than 50%) in their DNA and predominantly rod-shaped (bacillar) morphology, although some members are spherical (cocci). The name derives from the Latin word for "rod," reflecting the shape of its , Bacillus. This class encompasses a wide diversity of aerobic, microaerophilic, or facultatively species, many of which are capable of forming endospores that enable survival under extreme environmental conditions such as heat, , and radiation. Bacilli are ubiquitous in terrestrial and aquatic environments, including soil, water, air, and as commensals or pathogens in plants, animals, and humans. Taxonomically, Bacilli is defined based on 16S rRNA gene sequence analyses and includes at least two major orders: (the type order) and Lactobacillales. The order comprises genera such as the endospore-forming and non-endospore-forming and , with the latter including coccal forms despite the class's bacillar namesake. In contrast, Lactobacillales consists mainly of non-spore-forming, acid-tolerant lactic acid bacteria (LAB) like , , and , which are often associated with processes and host microbiomes. Recent genomic studies have revealed further within the class, including novel orders such as Erysipelotrichales and Mycoplasmatales, some of which exhibit reductive leading to loss of sporulation or even in their cell walls, adapting them to specialized niches like animal guts. This phylogenetic grouping highlights the class's evolutionary adaptability, with members spanning free-living to obligate parasites. Ecologically, Bacilli play crucial roles in nutrient cycling, , and symbiotic relationships. Species in , such as Bacillus subtilis, are key decomposers in soil, breaking down organic matter and promoting plant growth through production of plant growth-promoting rhizobacteria (PGPR) compounds like auxins and siderophores. LAB in Lactobacillales dominate fermented foods and the gastrointestinal of humans and animals, where they produce to inhibit pathogens and aid digestion. However, certain Bacilli are opportunistic or true pathogens; for instance, Bacillus anthracis causes , a zoonotic affecting and humans, while Listeria monocytogenes leads to , particularly dangerous for pregnant individuals and immunocompromised people. Streptococci in Lactobacillales are responsible for infections ranging from to severe invasive diseases like . In medicine and industry, Bacilli are invaluable for their biotechnological potential. Non-pathogenic strains of Bacillus species serve as probiotics to modulate gut health and enhance immune responses, while Bacillus thuringiensis produces insecticidal proteins used in biological pest control. LAB are essential in food preservation, producing yogurt, cheese, and sauerkraut through fermentation, and some synthesize bacteriocins as natural preservatives. Industrially, enzymes like subtilisin from B. subtilis are widely used in detergents and pharmaceuticals, underscoring the class's economic significance. Despite these benefits, antibiotic resistance in pathogenic Bacilli, such as methicillin-resistant Staphylococcus aureus (MRSA), poses ongoing public health challenges.

Terminology

Definition

Bacilli is a class of bacteria within the phylum ( Firmicutes), encompassing primarily Gram-positive characterized by their rod-shaped and low mol% G+C content (typically less than 50%). The class was formally defined by Ludwig et al. in 2010 based on phylogenetic analyses of 16S rRNA gene sequences, establishing it as a monophyletic group distinct from other Firmicutes classes like . This classification reflects the evolutionary relationships among endospore-forming and non-spore-forming lineages within the phylum. The class Bacilli includes major orders such as , which comprises endospore-forming rods and cocci such as those in the genus , and (also known as Lactobacillus order), consisting mainly of non-spore-forming like and , among others. Key diagnostic features of Bacilli include a Gram-positive structure and low mol% G+C content in their DNA, which distinguishes them from higher G+C Gram-positive phyla like . The name "Bacilli" derives from the Latin word , meaning "small rod," highlighting the predominant rod-shaped (bacilliform) of many members in this class, though some taxa exhibit coccal forms. This underscores the historical focus on cellular shape in early , while the modern definition emphasizes molecular and phylogenetic criteria.

Ambiguity with Shape and Other Taxa

The term "" (plural: bacilli) is frequently used in to describe the -shaped of bacterial cells, regardless of their taxonomic affiliation, leading to widespread when referring to the specific within the . This morphological descriptor applies to elongated, cylindrical forms observed across diverse bacterial groups, including both Gram-positive and Gram-negative species, and predates modern phylogenetic classifications based on 16S rRNA sequencing. For instance, , a Gram-negative commonly classified as a bacillus due to its shape, belongs to the in the , not the . This confusion is compounded by the historical origins of the terminology, where "bacilli" was first employed by Ferdinand Cohn in his 1872 treatise on bacteria to denote rod-like forms, long before the establishment of formal taxonomic hierarchies in bacteriology. Cohn's classification divided bacteria into four morphological groups—Sphaerobacteria (cocci), Microbacteria (short rods), Desmobacteria (filaments), and Spirobacteria (spirals)—with "bacillus" specifically introduced as a genus name for certain rod-shaped, spore-forming species like Bacillus subtilis, emphasizing form over genetic relatedness. This early emphasis on morphology as a classificatory criterion has persisted in common usage, often overshadowing the class Bacilli's broader phylogenetic scope, which encompasses not only rods but also cocci such as those in the genera Staphylococcus and Streptococcus. Further ambiguity arises from the overlap between the genus Bacillus—a group of primarily Gram-positive, endospore-forming rods within the order Bacillales of class Bacilli—and the class itself, leading to erroneous generalizations that equate all bacilli with members of this genus. For example, Gram-negative rods like Pseudomonas species, which are also termed bacilli morphologically, are classified in the class Gammaproteobacteria of the phylum Pseudomonadota, illustrating how the term transcends phylum boundaries. Such distinctions are critical in clinical and research contexts to avoid misidentification, as the class Bacilli is defined by shared genetic and physiological traits rather than shape alone.

Taxonomy and Phylogeny

Historical Development

The classification of Bacilli began in the mid-19th century with Ferdinand Cohn's foundational work on bacterial taxonomy, where he included rod-shaped bacteria, including what would later be recognized as Bacilli, within the group Schizomycetes, emphasizing their fission-based reproduction and morphological diversity. Cohn's 1875 publication marked an early attempt to systematize bacteria based on observable traits like shape and arrangement, laying the groundwork for subsequent refinements without yet distinguishing spore-forming capabilities. By the early , classifications evolved to incorporate physiological traits, as seen in Orla-Jensen's system, which refined Cohn's framework by separating spore-forming rods (assigned to ) from non-spore-formers, highlighting production as a key phenotypic marker for aerobic, . This approach initially relied heavily on formation and metabolic products like to delineate groups, with lactic acid production linking certain rods to fermentative processes, though genetic data was absent at the time. These phenotypic criteria dominated until molecular methods emerged, allowing for more precise groupings. In the late 20th century, broader taxonomic shifts occurred with Gibbons and Murray's 1978 proposal of the division Firmicutes to encompass all Gram-positive bacteria with low G+C content, initially positioning Bacilli as a class that included groups resembling Clostridia based on cell wall and spore traits. A pivotal milestone came in the 1980s, when 16S rRNA sequencing enabled the separation of Bacillales (primarily aerobic spore-formers) from Lactobacillales (lactic acid producers, mostly non-spore-forming), refining the class Bacilli to reflect phylogenetic relationships over purely morphological ones. This molecular approach supplanted earlier reliance on phenotypic traits, establishing Bacilli within Firmicutes as a distinct class by the 1990s. More recently, a 2021 reclassification proposed renaming Firmicutes to Bacillota for nomenclatural consistency, though it sparked debate and remains controversial among microbiologists.

Current Classification

The class Bacilli belongs to the phylum (previously designated Firmicutes), a phylum of predominantly characterized by low G+C content in their DNA. In contemporary taxonomy, Bacilli encompasses numerous orders (over 10 as of 2024 per LPSN), including the prominent and Lactobacillales, reflecting distinct phylogenetic clusters based on 16S rRNA gene sequences and whole-genome analyses. The order encompasses families such as (type family), , Paenibacillaceae, and Listeriaceae, while includes (type family), Streptococcaceae, Leuconostocaceae, and Enterococcaceae, among others. Additional orders include Erysipelotrichales and Mycoplasmatales, highlighting recent genomic-driven expansions. As documented in the List of Prokaryotic names with Standing in (LPSN) based on 2024 updates, the class comprises more than 10 orders, more than 50 families, and over 300 genera, with ongoing expansions driven by genomic data. Prominent genera within include (rod-shaped endospore formers) and (cocci in clusters), whereas Lactobacillales features (non-spore-forming rods) and (cocci in chains). Post-2010 taxonomic developments have integrated numerous novel genera from environmental isolates into Bacilli, exemplified by phylogenomic reclassifications within , such as the elevation of environmental Bacillus-like taxa into genera like and related groups.

Phylogenetic Analysis

Phylogenetic analyses of the Bacilli class primarily rely on molecular methods, including 16S rRNA gene sequencing and whole-genome-based approaches. The All-Species Living Tree Project (LTP) utilizes curated 16S rRNA sequences from type strains to construct phylogenetic trees, with the most recent updates incorporating alignments up to 2020 and ongoing integrations into SILVA databases for prokaryotic taxonomy. Complementing this, the Genome Taxonomy Database (GTDB) employs whole-genome sequences analyzed via concatenated alignments of 120 universal bacterial marker proteins to infer robust phylogenies, as implemented in release R10-RS226 from April 2025. These methods enable rank-normalized classifications that prioritize monophyly and genome similarity metrics like average nucleotide identity (ANI). Key findings from these analyses confirm that Bacilli forms a monophyletic within the phylum Firmicutes (now termed in GTDB), encompassing orders such as and Lactobacillales. Within this structure, occupies a basal position relative to Lactobacillales in phylogenomic trees derived from marker protein concatenates, reflecting deeper divergence among aerobic, spore-forming lineages. Bacilli shares a close evolutionary relationship with the class , another major Firmicutes group, but is distinguished by higher genomic G+C content (typically 35-55 mol%) compared to the lower values (20-45 mol%) prevalent in , correlating with differences in metabolic versatility and environmental adaptations. Discrepancies arise between GTDB and NCBI classifications, particularly in Lactobacillales, where GTDB has reclassified numerous genera based on phylogenomic coherence; for instance, the 2020 reorganization of the family split the polyphyletic genus into 25 novel genera to resolve non-monophyletic groupings unsupported by whole-genome data. In contrast, NCBI retains broader, phenotype-influenced groupings, leading to inconsistencies in species assignments for over 300 . Evolutionary insights highlight the role of (HGT) in shaping Bacilli genomes, especially for spore-forming genes; comparative analyses show that core sporulation regulators like spo0A are vertically inherited, but accessory genes involved in coat assembly exhibit HGT signatures, facilitating adaptation to diverse niches. The divergence of Bacilli from other Firmicutes lineages is estimated at 2-3 billion years ago, aligning with the emergence of formation near the base of the phylum during the .

Morphology and Physiology

Cellular Structure

Members of the class Bacilli are predominantly Gram-positive bacteria with diverse morphologies, typically rod-shaped (bacilliform) cells measuring 0.5–1.0 μm in width and 1.0–4.0 μm in length, though some lineages form cocci. For instance, genera like Bacillus and Lactobacillus exhibit rod forms, while Staphylococcus consists of cocci. These cells often arrange in pairs (diplobacilli) or chains (streptobacilli), depending on the species and growth conditions. The cell wall structure is a defining feature for most Bacilli, consisting of a thick layer that can comprise up to 90% of the wall's composition, conferring rigidity and Gram-positive staining. However, certain derived orders, such as Mycoplasmatales, have undergone reductive evolution, resulting in the loss of and s entirely; these wall-less are adapted to specialized niches like animal hosts and cannot undergo traditional Gram staining. Teichoic acids, anionic polymers of glycerol phosphate or phosphate, are covalently linked to the in wall-bearing species, aiding in cation and cell wall stability. Certain species, such as , produce a protective capsule composed of . Many rod-shaped Bacilli are motile, equipped with peritrichous flagella distributed over the cell surface to facilitate swimming in aqueous environments. In the order , a key ultrastructural adaptation is formation, producing one dormant per cell that resists heat, , and chemicals; these spores contain dipicolinic acid, which binds calcium to stabilize the core and enhance thermal resistance. In contrast, Lactobacillales lack this capability, as do many other orders including those with reductive evolution such as Mycoplasmatales.

Metabolic and Growth Characteristics

Bacilli display a range of metabolic strategies reflecting the physiological diversity within the class, particularly between its two primary orders, and Lactobacillales. Members of are predominantly aerobic or facultatively anaerobic, relying on respiratory to generate energy, and most produce to decompose into water and oxygen, aiding in management. In contrast, Lactobacillales are typically facultative anaerobes that favor fermentative pathways, with many species classified as that convert sugars into via homolactic or heterolactic , often lacking activity. This metabolic versatility enables Bacilli to adapt to varying oxygen availability and nutrient conditions in their environments. Growth characteristics of Bacilli are influenced by environmental factors such as temperature and , with optimal ranges varying by but generally centered around mesophilic conditions. Most Bacilli thrive at temperatures between 20°C and 40°C and neutral values of 6 to 7, though extremes exist; for instance, psychrotolerant strains can grow below 10°C, while thermophilic members like Geobacillus achieve optimal growth up to 70°C. The formation of endospores in many further enhances survival, allowing metabolic dormancy and resistance to harsh conditions like extreme heat, , or until favorable growth resumes. Nutritionally, Bacilli are predominantly heterotrophic, deriving energy from organic compounds such as sugars, , and proteins through chemoorganotrophic metabolism. Certain genera, including within , possess the capability for biological , utilizing enzymes to convert atmospheric N₂ into for assimilation, which supports growth in nitrogen-limited settings. Regarding antibiotic interactions, most Bacilli with intact Gram-positive cell walls exhibit sensitivity to penicillin due to the absence of an outer membrane barrier, allowing the antibiotic to target penicillin-binding proteins effectively. Wall-less lineages such as those in Mycoplasmatales, however, lack peptidoglycan and are inherently resistant to β-lactam antibiotics. Resistance emerges in some wall-bearing strains through the production of β-lactamases, enzymes that hydrolyze the β-lactam ring of penicillin, with examples including class D β-lactamases identified in species like Bacillus subtilis and Bacillus pumilus.

Ecology and Distribution

Habitats and Environments

Bacilli, as a class of primarily , exhibit a ubiquitous distribution across diverse ecosystems, including , , and air. In terrestrial environments, they are particularly prevalent in s, where genera such as often dominate rhizospheres, the nutrient-rich zones surrounding roots. This prevalence stems from their ability to thrive in organic-rich substrates like decomposing material. Aquatic systems also harbor significant populations, with isolates recovered from both sediments and freshwater bodies; some form clades adapted to varied salinities. Aerial dispersal further contributes to their global spread, allowing colonization of remote or transient environments. Extreme environments host specialized Bacilli, demonstrating their remarkable environmental tolerance. Thermotolerant species, such as certain Bacillus strains, inhabit hot springs with temperatures exceeding 60°C, where they contribute to microbial communities shaped by high thermal gradients. In acidic soils, populations persist at pH levels as low as 4-5, with densities ranging from 5 × 10^4 to 3 × 10^6 cells per gram in regions like coastal , . These adaptations enable survival in otherwise hostile conditions, including oligotrophic waters and desiccated lagoons. Neutral to alkaline soils support the highest abundances of Bacilli. and freshwater isolates similarly show densities in the 10^5 to 10^7 CFU per liter range in sediments. Key physiological adaptations underpin Bacilli's persistence in challenging habitats. Spore formation, a hallmark trait, confers resistance to , scarcity, and oligotrophic conditions, as seen in species like Bacillus coahuilensis from arid lagoons, where endospores endure prolonged . production on surfaces enhances adhesion and protection in fluctuating environments, such as air-water interfaces or aggregates. These mechanisms allow low-level persistence in biodiversity hotspots like animal gut microbiomes, where Bacilli constitute minor components (often <1% of total ), and plant endospheres, where endophytic strains contribute to internal without dominating communities. Recent studies as of 2024 have noted shifts in Bacilli communities in response to environmental changes, such as warming in aquatic habitats. Overall, these traits facilitate Bacilli's role as resilient opportunists in global microbial ecosystems.

Symbiotic and Pathogenic Interactions

Bacilli exhibit diverse symbiotic relationships, particularly mutualistic associations that benefit both the bacteria and their hosts. In the human gut microbiome, species of , such as L. rhamnosus and L. acidophilus, form symbiotic partnerships by fermenting carbohydrates and producing , which aids host digestion, enhances nutrient absorption, and maintains intestinal barrier integrity against pathogens. These bacteria also modulate the , promoting tolerance and reducing , thereby supporting overall gut . In plant-associated environments, certain Bacillus species, including B. subtilis and B. velezensis, engage in mutualistic with , promoting growth through mechanisms like solubilization. These secrete organic acids and enzymes that convert insoluble into bioavailable forms, enhancing uptake and improving yields in nutrient-poor soils. This interaction not only sustains bacterial populations via root exudates but also indirectly benefits the host by increasing to stresses. Many Bacilli are pathogenic, causing infections in humans and animals through opportunistic or primary mechanisms. , an opportunistic within the Bacilli class, commonly infects compromised skin sites such as wounds, leading to conditions like or abscesses by colonizing and proliferating in damaged tissues. In contrast, acts as a true , primarily causing acute (strep throat) by invading the oropharyngeal mucosa and eliciting inflammatory responses. Pathogenicity in Bacilli often relies on virulence factors that facilitate host invasion and immune evasion. For instance, S. pyogenes produces toxins such as streptolysins O and S, which lyse host cells, disrupt membranes, and promote tissue damage to aid bacterial spread. Invasins like lyase enable tissue penetration, while the capsule shields from , allowing evasion of innate immunity. Transmission of pathogenic Bacilli typically occurs from environmental reservoirs, such as or contaminated surfaces, to susceptible hosts via multiple routes. S. aureus spreads through direct contact with infected skin or fomites, as well as foodborne contamination leading to staphylococcal food poisoning. S. pyogenes is primarily transmitted via respiratory aerosols from coughing or sneezing, facilitating person-to-person spread in close-contact settings. These pathways underscore the importance of in preventing infections from ubiquitous Bacilli reservoirs.

Significance and Applications

Pathogenic Members

is a spore-forming bacterium responsible for , a zoonotic that manifests in cutaneous, inhalational, and gastrointestinal forms depending on the route of . Cutaneous , the most common form, results from spore entry through skin breaks, leading to localized lesions that can progress to systemic infection if untreated. Inhalational occurs via aerosolized spores, causing severe respiratory distress and high mortality without prompt intervention, while gastrointestinal arises from ingesting contaminated meat, resulting in and bloody . is primarily spore-mediated, with spores persisting in and infecting humans through contact with infected animals or their products. Vaccination efforts include the Sterne strain for , which provides protective immunity against virulent strains, and the vaccine adsorbed (AVA) for at-risk humans, administered as a series of doses. Staphylococcus aureus, particularly methicillin-resistant strains (MRSA), causes a spectrum of infections ranging from skin and abscesses to severe and . MRSA infections often begin as localized skin lesions but can disseminate, leading to with mortality rates exceeding 20% in hospitalized patients. Many S. aureus infections are toxin-mediated, with (TSST-1) implicated in , which triggers massive release and multi-organ failure. TSST-1-producing strains are associated with approximately 50% of non-menstrual toxic shock cases and nearly all menstrual cases. Streptococcus pyogenes, known as group A Streptococcus (GAS), is a major cause of pyogenic infections including and , a rapidly progressive tissue-destroying condition. presents with fever, rash, and strawberry tongue following , while involves deep tissue necrosis and requires urgent surgical . Diagnosis of recent GAS infection often relies on (ASO) titers, which measure antibodies against streptolysin O and rise 1-3 weeks post-infection, aiding in confirming post-streptococcal complications like . Epidemiologically, anthrax reports approximately 2,000-20,000 human cases annually worldwide, predominantly in developing regions with exposure. As estimated in 2005, GAS diseases impose a global burden exceeding 18 million serious cases yearly, with over 500,000 deaths linked to invasive infections and sequelae like rheumatic heart disease. S. aureus contributes significantly to (AMR) threats, with MRSA bloodstream infections accounting for a substantial portion of the 1.27 million AMR-attributable deaths in 2019. Post-2020, AMR trends in these Bacilli pathogens have intensified due to disruptions, including increased MRSA resistance proportions in healthcare settings and rising GAS bacteremia incidence.

Biotechnological and Industrial Roles

Bacilli species, particularly those in the genera and , play pivotal roles in biotechnological and industrial applications due to their robust enzyme production, capabilities, and potential. These Gram-positive, spore-forming or are valued for their stability under diverse conditions, enabling scalable processes in food, pharmaceutical, and environmental sectors. In enzyme production, is a primary source of , an alkaline widely incorporated into laundry detergents for its ability to break down protein-based stains at elevated temperatures and levels. This enzyme's and activity in the presence of make it essential for modern detergent formulations, with industrial yields optimized through and techniques. Similarly, Geobacillus species, such as G. stearothermophilus, produce thermostable α-amylases used in hydrolysis for industries like , textiles, and biofuels, where the enzymes maintain activity above 60°C, facilitating high-temperature reactions that enhance efficiency and reduce contamination risks. Fermentation processes leverage species for producing fermented dairy and vegetable products, including , cheese, and , where they convert sugars into , lowering to preserve food and impart characteristic flavors and textures. For instance, Lactobacillus bulgaricus and are key in production, initiating rapid acidification that coagulates milk proteins. Additionally, serves as a in supplements and fortified foods, promoting gut health by balancing intestinal , inhibiting pathogens, and aiding digestion, with clinical evidence supporting its role in alleviating and . In , species demonstrate efficacy in degrading pesticides, such as and , through enzymatic hydrolysis that mineralizes these compounds into less toxic byproducts, offering a sustainable alternative to chemical treatments in contaminated soils and waters. strains, for example, have been shown to achieve over 80% degradation of within days under aerobic conditions. These also facilitate sequestration via and mechanisms, where extracellular polymeric substances bind ions like lead, , and , reducing their and in polluted environments; B. cereus and B. subtilis exhibit removal efficiencies up to 90% for multiple metals in lab-scale trials. Pharmaceutical applications of Bacilli include antibiotic production, with Bacillus licheniformis serving as the main producer of bacitracin, a effective against and used topically for wound infections and in to prevent necrotic . Fermentation optimization has increased yields to over 700 IU/mL through targeting precursor pathways. Furthermore, Bacillus subtilis spores function as safe, non-pathogenic vaccine vectors, displaying antigens on their surface for mucosal delivery; this platform has been explored for vaccines against and , eliciting strong immune responses in animal models due to the spores' stability and adjuvanticity.