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In bacteriology, gram-positive bacteria are
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
that give a positive result in the Gram stain test, which is traditionally used to quickly classify bacteria into two broad categories according to their type of
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mec ...
. Gram-positive bacteria take up the
crystal violet Crystal violet or gentian violet, also known as methyl violet 10B or hexamethyl pararosaniline chloride, is a triarylmethane dye used as a histological stain and in Gram's method of classifying bacteria. Crystal violet has antibacterial, antif ...
stain used in the test, and then appear to be purple-coloured when seen through an
optical microscope The optical microscope, also referred to as a light microscope, is a type of microscope that commonly uses visible light and a system of lenses to generate magnified images of small objects. Optical microscopes are the oldest design of micro ...
. This is because the thick peptidoglycan layer in the bacterial cell wall retains the
stain A stain is a discoloration that can be clearly distinguished from the surface, material, or medium it is found upon. They are caused by the chemical or physical interaction of two dissimilar materials. Accidental staining may make materials ap ...
after it is washed away from the rest of the sample, in the decolorization stage of the test. Conversely, gram-negative bacteria cannot retain the violet stain after the decolorization step; alcohol used in this stage degrades the outer membrane of gram-negative cells, making the cell wall more porous and incapable of retaining the crystal violet stain. Their peptidoglycan layer is much thinner and sandwiched between an
inner cell membrane The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the cytoplasm, interior of all Cell (biology), cells from th ...
and a bacterial outer membrane, causing them to take up the
counterstain A counterstain is a stain with colour contrasting to the principal stain, making the stained structure easily visible using a microscope. Examples include the malachite green counterstain to the fuchsine stain in the Gimenez staining technique ...
( safranin or fuchsine) and appear red or pink. Despite their thicker peptidoglycan layer, gram-positive bacteria are more receptive to certain
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mec ...
targeting
antibiotics An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important type of antibacterial agent for fighting bacterial infections, and antibiotic medications are widely used in the treatment and preventio ...
than gram-negative bacteria, due to the absence of the outer membrane.


Characteristics

In general, the following characteristics are present in gram-positive bacteria: # Cytoplasmic lipid membrane # Thick peptidoglycan layer #
Teichoic acids Teichoic acids (''cf.'' Greek τεῖχος, ''teīkhos'', "wall", to be specific a fortification wall, as opposed to τοῖχος, ''toīkhos'', a regular wall) are bacterial copolymers of glycerol phosphate or ribitol phosphate and carboh ...
and lipoids are present, forming
lipoteichoic acid Lipoteichoic acid (LTA) is a major constituent of the cell wall of gram-positive bacteria. These organisms have an inner (or cytoplasmic) membrane and, external to it, a thick (up to 80 nanometer) peptidoglycan layer. The structure of LTA varies ...
s, which serve as chelating agents, and also for certain types of adherence. # Peptidoglycan chains are cross-linked to form rigid cell walls by a bacterial enzyme
DD-transpeptidase DD-transpeptidase (, ''DD-peptidase'', ''DD-transpeptidase'', ''DD-carboxypeptidase'', ''D-alanyl-D-alanine carboxypeptidase'', ''D-alanyl-D-alanine-cleaving-peptidase'', ''D-alanine carboxypeptidase'', ''D-alanyl carboxypeptidase'', and ''serine-t ...
. # A much smaller volume of periplasm than that in gram-negative bacteria. Only some species have a capsule, usually consisting of polysaccharides. Also, only some species are flagellates, and when they do have flagella, have only two basal body rings to support them, whereas gram-negative have four. Both gram-positive and gram-negative bacteria commonly have a surface layer called an S-layer. In gram-positive bacteria, the S-layer is attached to the peptidoglycan layer. Gram-negative bacteria's S-layer is attached directly to the outer membrane. Specific to gram-positive bacteria is the presence of teichoic acids in the cell wall. Some of these are lipoteichoic acids, which have a lipid component in the cell membrane that can assist in anchoring the peptidoglycan.


Classification

Along with cell shape, Gram staining is a rapid method used to differentiate bacterial species. Such staining, together with growth requirement and antibiotic susceptibility testing, and other macroscopic and physiologic tests, forms the full basis for classification and subdivision of the bacteria (e.g., see figure and pre-1990 versions of ''
Bergey's Manual of Systematic Bacteriology ''Bergey's Manual of Systematic Bacteriology'' is the main resource for determining the identity of prokaryotic organisms, emphasizing bacterial species, using every characterizing aspect. The manual was published subsequent to the ''Bergey's Man ...
'').
Historically History (derived ) is the systematic study and the documentation of the human activity. The time period of event before the invention of writing systems is considered prehistory. "History" is an umbrella term comprising past events as well ...
, the kingdom Monera was divided into four divisions based primarily on Gram staining: Bacillota (positive in staining), Gracilicutes (negative in staining), Mollicutes (neutral in staining) and Mendocutes (variable in staining). Based on 16S ribosomal RNA phylogenetic studies of the late microbiologist Carl Woese and collaborators and colleagues at the
University of Illinois The University of Illinois Urbana-Champaign (U of I, Illinois, University of Illinois, or UIUC) is a public land-grant research university in Illinois in the twin cities of Champaign and Urbana. It is the flagship institution of the Uni ...
, the
monophyly In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic ...
of the gram-positive bacteria was challenged, with major implications for the therapeutic and general study of these organisms. Based on molecular studies of the 16S sequences, Woese recognised twelve bacterial phyla. Two of these were gram-positive and were divided on the proportion of the
guanine Guanine () ( symbol G or Gua) is one of the four main nucleobases found in the nucleic acids DNA and RNA, the others being adenine, cytosine, and thymine ( uracil in RNA). In DNA, guanine is paired with cytosine. The guanine nucleoside is ...
and
cytosine Cytosine () ( symbol C or Cyt) is one of the four nucleobases found in DNA and RNA, along with adenine, guanine, and thymine ( uracil in RNA). It is a pyrimidine derivative, with a heterocyclic aromatic ring and two substituents attached ( ...
content in their DNA. The high G + C phylum was made up of the Actinobacteria and the low G + C phylum contained the Firmicutes. The Actinomycetota include the '' Corynebacterium'', '' Mycobacterium'', '' Nocardia'' and ''
Streptomyces ''Streptomyces'' is the largest genus of Actinomycetota and the type genus of the family Streptomycetaceae. Over 500 species of ''Streptomyces'' bacteria have been described. As with the other Actinomycetota, streptomycetes are gram-positiv ...
'' genera. The (low G + C) Bacillota, have a 45–60% GC content, but this is lower than that of the Actinomycetota.


Importance of the outer cell membrane in bacterial classification

Although bacteria are traditionally divided into two main groups, gram-positive and gram-negative, based on their Gram stain retention property, this classification system is ambiguous as it refers to three distinct aspects (staining result, envelope organization, taxonomic group), which do not necessarily coalesce for some bacterial species. The gram-positive and gram-negative staining response is also not a reliable characteristic as these two kinds of bacteria do not form phylogenetic coherent groups. However, although Gram staining response is an empirical criterion, its basis lies in the marked differences in the ultrastructure and chemical composition of the bacterial cell wall, marked by the absence or presence of an outer lipid membrane. All gram-positive bacteria are bounded by a single-unit lipid membrane, and, in general, they contain a thick layer (20–80 nm) of peptidoglycan responsible for retaining the Gram stain. A number of other bacteria—that are bounded by a single membrane, but stain gram-negative due to either lack of the peptidoglycan layer, as in the mycoplasmas, or their inability to retain the Gram stain because of their cell wall composition—also show close relationship to the Gram-positive bacteria. For the bacterial cells bounded by a single cell membrane, the term ''monoderm bacteria'' has been proposed. In contrast to gram-positive bacteria, all typical gram-negative bacteria are bounded by a cytoplasmic membrane and an outer cell membrane; they contain only a thin layer of peptidoglycan (2–3 nm) between these membranes. The presence of inner and outer cell membranes defines a new compartment in these cells: the periplasmic space or the periplasmic compartment. These bacteria have been designated as
diderm bacteria Gram-negative bacteria are bacteria that do not retain the crystal violet stain used in the Gram staining method of bacterial differentiation. They are characterized by their cell envelopes, which are composed of a thin peptidoglycan cell wall ...
. The distinction between the monoderm and diderm bacteria is supported by conserved signature indels in a number of important proteins (viz. DnaK, GroEL). Of these two structurally distinct groups of bacteria, monoderms are indicated to be ancestral. Based upon a number of observations including that the gram-positive bacteria are the major producers of antibiotics and that, in general, gram-negative bacteria are resistant to them, it has been proposed that the outer cell membrane in gram-negative bacteria (diderms) has evolved as a protective mechanism against
antibiotic An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important type of antibacterial agent for fighting bacterial infections, and antibiotic medications are widely used in the treatment and prevention ...
selection pressure. Some bacteria, such as ''
Deinococcus ''Deinococcus'' (from the el, δεινός, ''deinos'', "dreadful, strange" and κόκκος, ''kókkos'', "granule") is in the monotypic family Deinococcaceae, and one genus of three in the order Deinococcales of the bacterial phylum '' Deinoco ...
'', which stain gram-positive due to the presence of a thick peptidoglycan layer and also possess an outer cell membrane are suggested as intermediates in the transition between monoderm (gram-positive) and diderm (gram-negative) bacteria. The diderm bacteria can also be further differentiated between simple diderms lacking lipopolysaccharide, the archetypical diderm bacteria where the outer cell membrane contains lipopolysaccharide, and the diderm bacteria where outer cell membrane is made up of mycolic acid.


Exceptions

In general, gram-positive bacteria are monoderms and have a single lipid bilayer whereas gram-negative bacteria are diderms and have two bilayers. Some taxa lack peptidoglycan (such as the class Mollicutes, some members of the Rickettsiales, and the insect-endosymbionts of the Enterobacteriales) and are gram-variable. This, however, does not always hold true. The Deinococcota have gram-positive stains, although they are structurally similar to gram-negative bacteria with two layers. The
Chloroflexota The Chloroflexota are a phylum of bacteria containing isolates with a diversity of phenotypes, including members that are aerobic thermophiles, which use oxygen and grow well in high temperatures; anoxygenic phototrophs, which use light for p ...
have a single layer, yet (with some exceptions) stain negative. Two related phyla to the Chloroflexi, the
TM7 Saccharibacteria, formerly known as ''TM7'', is a major bacterial lineage. It was discovered through 16S rRNA sequencing . TM7x from the human oral cavity was cultivated and revealed that TM7x is an extremely small coccus (200-300 nm) an ...
clade and the Ktedonobacteria, are also monoderms. Some Bacillota species are not gram-positive. These belong to the class Mollicutes (alternatively considered a class of the phylum Mycoplasmatota), which lack peptidoglycan ( gram-indeterminate), and the class Negativicutes, which includes ''
Selenomonas Members of the genus ''Selenomonas'' (motile crescent-shaped bacteria in general) are referred to trivially as selenomonads. The genus ''Selenomonas'' constitutes a group of motile crescent-shaped bacteria and includes species living in the gast ...
'' and stain gram-negative. Additionally, a number of bacterial taxa (viz. Negativicutes, Fusobacteriota, Synergistota, and Elusimicrobiota) that are either part of the phylum Bacillota or branch in its proximity are found to possess a diderm cell structure. However, a conserved signature indel (CSI) in the HSP60 ( GroEL) protein distinguishes all traditional phyla of gram-negative bacteria (e.g., Pseudomonadota, Aquificota,
Chlamydiota The Chlamydiota (synonym Chlamydiae) are a bacterial phylum and class whose members are remarkably diverse, including pathogens of humans and animals, symbionts of ubiquitous protozoa, and marine sediment forms not yet well understood. All of ...
,
Bacteroidota The phylum Bacteroidota (synonym Bacteroidetes) is composed of three large classes of Gram-negative, nonsporeforming, anaerobic or aerobic, and rod-shaped bacteria that are widely distributed in the environment, including in soil, sediments, and ...
,
Chlorobiota The green sulfur bacteria are a phylum of obligately anaerobic photoautotrophic bacteria that metabolize sulfur. Green sulfur bacteria are nonmotile (except ''Chloroherpeton thalassium'', which may glide) and capable of anoxygenic photosynthes ...
, "
Cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, bl ...
", Fibrobacterota, Verrucomicrobiota, Planctomycetota, Spirochaetota,
Acidobacteriota Acidobacteriota is a phylum of Gram-negative bacteria. Its members are physiologically diverse and ubiquitous, especially in soils, but are under-represented in culture. Description Members of this phylum are physiologically diverse, and can be ...
, etc.) from these other atypical diderm bacteria, as well as other phyla of monoderm bacteria (e.g., Actinomycetota, Bacillota, Thermotogota,
Chloroflexota The Chloroflexota are a phylum of bacteria containing isolates with a diversity of phenotypes, including members that are aerobic thermophiles, which use oxygen and grow well in high temperatures; anoxygenic phototrophs, which use light for p ...
, etc.). The presence of this CSI in all sequenced species of conventional LPS (
lipopolysaccharide Lipopolysaccharides (LPS) are large molecules consisting of a lipid and a polysaccharide that are bacterial toxins. They are composed of an O- antigen, an outer core, and an inner core all joined by a covalent bond, and are found in the out ...
)-containing gram-negative bacterial phyla provides evidence that these phyla of bacteria form a monophyletic clade and that no loss of the outer membrane from any species from this group has occurred.


Pathogenicity

In the classical sense, six gram-positive genera are typically pathogenic in humans. Two of these, '' Streptococcus'' and '' Staphylococcus'', are cocci (sphere-shaped). The remaining organisms are bacilli (rod-shaped) and can be subdivided based on their ability to form
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...
s. The non-spore formers are '' Corynebacterium'' and '' Listeria'' (a coccobacillus), whereas '' Bacillus'' and ''
Clostridium ''Clostridium'' is a genus of anaerobic, Gram-positive bacteria. Species of ''Clostridium'' inhabit soils and the intestinal tract of animals, including humans. This genus includes several significant human pathogens, including the causative a ...
'' produce spores. The spore-forming bacteria can again be divided based on their respiration: ''Bacillus'' is a
facultative anaerobe A facultative anaerobic organism is an organism that makes ATP by aerobic respiration if oxygen is present, but is capable of switching to fermentation if oxygen is absent. Some examples of facultatively anaerobic bacteria are '' Staphylococ ...
, while ''Clostridium'' is an obligate anaerobe. Also, ''Rathybacter'', ''Leifsonia'', and ''Clavibacter'' are three gram-positive genera that cause plant disease. Gram-positive bacteria are capable of causing serious and sometimes fatal
infections An infection is the invasion of tissues by pathogens, their multiplication, and the reaction of host tissues to the infectious agent and the toxins they produce. An infectious disease, also known as a transmissible disease or communicable di ...
in newborn infants. Access provided by the University of Pittsburgh. Novel species of clinically relevant gram-positive bacteria also include ''
Catabacter hongkongensis ''Christensenella hongkongensis'' is a species of clinically relevant gram-positive coccobacilli, first isolated from patients in Hong Kong and Canada in 2006. Although the species remains relatively rare, it has a high mortality rate of up to ...
'', which is an emerging pathogen belonging to Bacillota.


Bacterial transformation

Transformation is one of three processes for horizontal gene transfer, in which exogenous genetic material passes from a donor bacterium to a recipient bacterium, the other two processes being
conjugation Conjugation or conjugate may refer to: Linguistics *Grammatical conjugation, the modification of a verb from its basic form * Emotive conjugation or Russell's conjugation, the use of loaded language Mathematics *Complex conjugation, the change ...
(transfer of genetic material between two bacterial cells in direct contact) and transduction (injection of donor bacterial DNA by a
bacteriophage A bacteriophage (), also known informally as a ''phage'' (), is a duplodnaviria virus that infects and replicates within bacteria and archaea. The term was derived from "bacteria" and the Greek φαγεῖν ('), meaning "to devour". Bac ...
virus into a recipient host bacterium). In transformation, the genetic material passes through the intervening medium, and uptake is completely dependent on the recipient bacterium. As of 2014 about 80 species of bacteria were known to be capable of transformation, about evenly divided between gram-positive and gram-negative bacteria; the number might be an overestimate since several of the reports are supported by single papers. Transformation among gram-positive bacteria has been studied in medically important species such as '' Streptococcus pneumoniae'', '' Streptococcus mutans'', '' Staphylococcus aureus'' and '' Streptococcus sanguinis'' and in gram-positive soil bacterium '' Bacillus subtilis, Bacillus cereus''.


Orthographic note

The adjectives ''Gram-positive'' and ''Gram-negative'' derive from the surname of Hans Christian Gram; as eponymous adjectives, their initial letter can be either capital ''G'' or lower-case ''g'', depending on which
style guide A style guide or manual of style is a set of standards for the writing, formatting, and design of documents. It is often called a style sheet, although that term also has multiple other meanings. The standards can be applied either for gene ...
(e.g., that of the
CDC The Centers for Disease Control and Prevention (CDC) is the national public health agency of the United States. It is a United States federal agency, under the Department of Health and Human Services, and is headquartered in Atlanta, Georg ...
), if any, governs the document being written. This is further explained at '' Gram staining § Orthographic note''.


References


External links

*
3D structures of proteins associated with plasma membrane of gram-positive bacteria

3D structures of proteins associated with outer membrane of gram-positive bacteria
{{Authority control Staining Bacteriology