cell Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery ...

biological Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary in ...

archaea Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaebac ...

eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...

biochemical Biochemistry or biological chemistry is the study of chemical processes within and relating to living organisms. A sub-discipline of both chemistry and biology, biochemistry may be divided into three fields: structural biology, enzymology an ...

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 among ...

morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines * Morphology (archaeology), study of the shapes or forms of artifacts * Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies ...

coccus A coccus (plural cocci) is any bacterium or archaeon that has a spherical, ovoid, or generally round shape. Bacteria are categorized based on their shapes into three classes: cocci (spherical-shaped), bacillus (rod-shaped) and spiral ( of whi ...

bacillus ''Bacillus'' (Latin "stick") is a genus of Gram-positive, rod-shaped bacteria, a member of the phylum ''Bacillota'', with 266 named species. The term is also used to describe the shape (rod) of other so-shaped bacteria; and the plural ''Bacilli ...

coccobacillus A coccobacillus (plural coccobacilli), or bacilluscocco, is a type of bacterium with a shape intermediate between cocci (spherical bacteria) and bacilli (rod-shaped bacteria). Coccobacilli, then, are very short rods which may be mistaken for coc ...

spiral In mathematics, a spiral is a curve which emanates from a point, moving farther away as it revolves around the point. Helices Two major definitions of "spiral" in the American Heritage Dictionary are:filamentous (elongated) Cell shape is generally characteristic of a given bacterial species, but can vary depending on growth conditions. Some bacteria have complex life cycles involving the production of stalks and appendages (e.g. '' Caulobacter'') and some produce elaborate structures bearing reproductive spores (e.g. ''

Myxococcus ''Myxococcus'' is a genus of bacteria in the family Myxococcaceae. Myxococci are Gram-negative, spore-forming, chemoorganotrophic, obligate aerobes. They are elongated rods with rounded or tapered ends, and they are nonflagellated. The cells mov ...

'', ''

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-positive, ...

'').

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 among ...

generally form distinctive cell morphologies when examined by

light microscopy Microscopy is the technical field of using microscopes to view objects and areas of objects that cannot be seen with the naked eye (objects that are not within the resolution range of the normal eye). There are three well-known branches of micr ...

and distinct colony morphologies when grown on Petri plates. Perhaps the most obvious structural characteristic of

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 among ...

is (with some exceptions) their small size. For example, ''

Escherichia coli ''Escherichia coli'' (),Wells, J. C. (2000) Longman Pronunciation Dictionary. Harlow ngland Pearson Education Ltd. also known as ''E. coli'' (), is a Gram-negative, facultative anaerobic, rod-shaped, coliform bacterium of the genus ''Escher ...

'' cells, an "average" sized bacterium, are about 2 µm (

micrometre The micrometre ( international spelling as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American spelling), also commonly known as a micron, is a unit of length in the International System of Unit ...

s) long and 0.5 µm in diameter, with a cell volume of 0.6–0.7 μm³. This corresponds to a wet mass of about 1

picogram To help compare different orders of magnitude, the following lists describe various mass levels between 10−59 kg and 1052 kg. The least massive thing listed here is a graviton, and the most massive thing is the observable universe. ...

(pg), assuming that the cell consists mostly of water. The dry mass of a single cell can be estimated as 23% of the wet mass, amounting to 0.2 pg. About half of the dry mass of a bacterial cell consists of carbon, and also about half of it can be attributed to proteins. Therefore, a typical fully grown 1-liter culture of ''

Escherichia coli ''Escherichia coli'' (),Wells, J. C. (2000) Longman Pronunciation Dictionary. Harlow ngland Pearson Education Ltd. also known as ''E. coli'' (), is a Gram-negative, facultative anaerobic, rod-shaped, coliform bacterium of the genus ''Escher ...

'' (at an optical density of 1.0, corresponding to c. 10⁹ cells/ml) yields about 1 g wet cell mass. Small size is extremely important because it allows for a large

surface area-to-volume ratio The surface-area-to-volume ratio, also called the surface-to-volume ratio and variously denoted sa/vol or SA:V, is the amount of surface area per unit volume of an object or collection of objects. SA:V is an important concept in science and engin ...

which allows for rapid uptake and intracellular distribution of nutrients and excretion of wastes. At low surface area-to-volume ratios the diffusion of nutrients and waste products across the bacterial cell membrane limits the rate at which microbial metabolism can occur, making the cell less evolutionarily fit. The reason for the existence of large cells is unknown, although it is speculated that the increased cell volume is used primarily for storage of excess nutrients. Comparison of a typical bacterial cell and a typical human cell (assuming both cells are spheres) :

Cell wall

The

cell envelope The cell envelope comprises the inner cell membrane and the cell wall of a bacterium. In gram-negative bacteria an outer membrane is also included. This envelope is not present in the Mollicutes where the cell wall is absent. Bacterial cell env ...

is composed of the

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 interior of all cells from the outside environment ( ...

and the

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 mech ...

. As in other organisms, the

bacterial 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 mech ...

provides structural integrity to the cell. In

prokaryote A prokaryote () is a single-celled organism that lacks a nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Greek πρό (, 'before') and κάρυον (, 'nut' or 'kernel').Campbell, N. "Biology:Concepts & Connec ...

s, the primary function of the cell wall is to protect the cell from internal

turgor pressure Turgor pressure is the force within the cell that pushes the plasma membrane against the cell wall. It is also called ''hydrostatic pressure'', and is defined as the pressure in a fluid measured at a certain point within itself when at equilibri ...

caused by the much higher concentrations of proteins, and other molecules inside the cell compared to its external environment. The bacterial cell wall differs from that of all other organisms by the presence of

peptidoglycan Peptidoglycan or murein is a unique large macromolecule, a polysaccharide, consisting of sugars and amino acids that forms a mesh-like peptidoglycan layer outside the plasma membrane, the rigid cell wall (murein sacculus) characteristic of most ...

which is located immediately outside of the cell membrane. Peptidoglycan is made up of a polysaccharide backbone consisting of alternating

N-Acetylmuramic acid ''N''-Acetylmuramic acid (NAM or MurNAc) is an organic compound with the chemical formula . It is a monomer of peptidoglycan in most bacterial cell walls, which is built from alternating units of ''N''-acetylglucosamine (GlcNAc) and ''N''-acet ...

(NAM) and

N-acetylglucosamine ''N''-Acetylglucosamine (GlcNAc) is an amide derivative of the monosaccharide glucose. It is a secondary amide between glucosamine and acetic acid. It is significant in several biological systems. It is part of a biopolymer in the bacterial ...

(NAG) residues in equal amounts.

Peptidoglycan Peptidoglycan or murein is a unique large macromolecule, a polysaccharide, consisting of sugars and amino acids that forms a mesh-like peptidoglycan layer outside the plasma membrane, the rigid cell wall (murein sacculus) characteristic of most ...

is responsible for the rigidity of the bacterial cell wall, and for the determination of cell shape. It is relatively porous and is not considered to be a permeability barrier for small substrates. While all bacterial cell walls (with a few exceptions such as extracellular

parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...

s such as ''

Mycoplasma ''Mycoplasma'' is a genus of bacteria that, like the other members of the class ''Mollicutes'', lack a cell wall around their cell membranes. Peptidoglycan (murein) is absent. This characteristic makes them naturally resistant to antibiotics ...

'') contain peptidoglycan, not all cell walls have the same overall structures. Since the cell wall is required for bacterial survival, but is absent in some

eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...

s, several

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 prevention o ...

(notably the

penicillin Penicillins (P, PCN or PEN) are a group of β-lactam antibiotics originally obtained from ''Penicillium'' moulds, principally '' P. chrysogenum'' and '' P. rubens''. Most penicillins in clinical use are synthesised by P. chrysogenum using ...

s and

cephalosporin The cephalosporins (sg. ) are a class of β-lactam antibiotics originally derived from the fungus ''Acremonium'', which was previously known as ''Cephalosporium''. Together with cephamycins, they constitute a subgroup of β-lactam antibiotics ...

s) stop bacterial infections by interfering with cell wall synthesis, while having no effects on

human cells There are many different types of cells in the human body. Cells derived primarily from endoderm Exocrine secretory epithelial cells * Brunner's gland cell in duodenum (enzymes and alkaline mucus) *Insulated goblet cell of respiratory and ...

which have no cell wall, only a cell membrane. There are two main types of bacterial cell walls, those of

gram-positive bacteria In bacteriology, gram-positive bacteria are bacteria 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. Gram-positive bact ...

and those of

gram-negative 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 ...

, which are differentiated by their

Gram staining In microbiology and bacteriology, Gram stain (Gram staining or Gram's method), is a method of staining used to classify bacterial species into two large groups: gram-positive bacteria and gram-negative bacteria. The name comes from the Danish bac ...

characteristics. For both these types of bacteria, particles of approximately 2 nm can pass through the peptidoglycan. If the bacterial cell wall is entirely removed, it is called a protoplast while if it's partially removed, it is called a

spheroplast A spheroplast (or sphaeroplast in British usage) is a microbial cell from which the cell wall has been almost completely removed, as by the action of penicillin or lysozyme. According to some definitions, the term is used to describe Gram-negative ...

.

Beta-lactam antibiotic β-lactam antibiotics (beta-lactam antibiotics) are antibiotics that contain a beta-lactam ring in their chemical structure. This includes penicillin derivatives (penams), cephalosporins and cephamycins (cephems), monobactams, carbapenems and ...

s such as penicillin inhibit the formation of peptidoglycan cross-links in the bacterial cell wall. The enzyme

lysozyme Lysozyme (EC 3.2.1.17, muramidase, ''N''-acetylmuramide glycanhydrolase; systematic name peptidoglycan ''N''-acetylmuramoylhydrolase) is an antimicrobial enzyme produced by animals that forms part of the innate immune system. It is a glycoside ...

, found in human tears, also digests the cell wall of bacteria and is the body's main defense against eye infections.

The gram-positive cell wall

Gram-positive cell walls are thick and the

peptidoglycan Peptidoglycan or murein is a unique large macromolecule, a polysaccharide, consisting of sugars and amino acids that forms a mesh-like peptidoglycan layer outside the plasma membrane, the rigid cell wall (murein sacculus) characteristic of most ...

(also known as ''murein'') layer constitutes almost 95% of the cell wall in some gram-positive bacteria and as little as 5-10% of the cell wall in gram-negative bacteria. The 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 ...

dye and are stained purple. The cell wall of some gram-positive bacteria can be completely dissolved by

lysozyme Lysozyme (EC 3.2.1.17, muramidase, ''N''-acetylmuramide glycanhydrolase; systematic name peptidoglycan ''N''-acetylmuramoylhydrolase) is an antimicrobial enzyme produced by animals that forms part of the innate immune system. It is a glycoside ...

s which attacks the bonds between N-acetylmuramic acid and N-acetylglucosamine. In other gram-positive bacteria, such as ''

Staphylococcus aureus ''Staphylococcus aureus'' is a Gram-positive spherically shaped bacterium, a member of the Bacillota, and is a usual member of the microbiota of the body, frequently found in the upper respiratory tract and on the skin. It is often positive ...

'', the walls are resistant to the action of lysozymes. They have O-acetyl groups on carbon-6 of some muramic acid residues. The matrix substances in the walls of gram-positive bacteria may be polysaccharides or

teichoic acid 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 carbohydr ...

s. The latter are very widespread, but have been found only in gram-positive bacteria. There are two main types of teichoic acid: ribitol teichoic acids and glycerol teichoic

acid In computer science, ACID ( atomicity, consistency, isolation, durability) is a set of properties of database transactions intended to guarantee data validity despite errors, power failures, and other mishaps. In the context of databases, a sequ ...

s. The latter one is more widespread. These acids are polymers of

ribitol Ribitol, or adonitol, is a crystalline pentose alcohol (C5H12O5) formed by the reduction of ribose. It occurs naturally in the plant '' Adonis vernalis'' as well as in the cell walls of some Gram-positive bacteria, in the form of ribitol phosp ...

phosphate and

glycerol phosphate Glycerol phosphate may refer to: * Glycerol 1-phosphate * Glycerol 2-phosphate * Glycerol 3-phosphate ''sn''-Glycerol 3-phosphate is the organic ion with the formula HOCH2CH(OH)CH2OPO32-. It is one of three stereoisomers of the ester of dibasi ...

, respectively, and only located on the surface of many gram-positive bacteria. However, the exact function of teichoic acid is debated and not fully understood. A major component of the gram-positive cell wall is

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 be ...

. One of its purposes is providing an antigenic function. The lipid element is to be found in the membrane where its adhesive properties assist in its anchoring to the membrane.

The gram-negative cell wall

Gram-negative cell walls are much thinner than the gram-positive cell walls, and they contain a second plasma membrane superficial to their thin

peptidoglycan Peptidoglycan or murein is a unique large macromolecule, a polysaccharide, consisting of sugars and amino acids that forms a mesh-like peptidoglycan layer outside the plasma membrane, the rigid cell wall (murein sacculus) characteristic of most ...

layer, in turn adjacent to the

cytoplasmic 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 interior of all cells from the outside environment ( ...

. Gram-negative bacteria are stained as pink colour. The chemical structure of the outer membrane's

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 outer m ...

is often unique to specific bacterial sub-species and is responsible for many of the

antigen In immunology, an antigen (Ag) is a molecule or molecular structure or any foreign particulate matter or a pollen grain that can bind to a specific antibody or T-cell receptor. The presence of antigens in the body may trigger an immune response. ...

ic properties of these strains.

Plasma membrane

The

plasma 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 interior of all cells from the outside environment (t ...

or bacterial cytoplasmic membrane is composed of a

phospholipid bilayer The lipid bilayer (or phospholipid bilayer) is a thin polar membrane made of two layers of lipid molecules. These membranes are flat sheets that form a continuous barrier around all cells. The cell membranes of almost all organisms and many vir ...

and thus has all of the general functions of a

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 interior of all cells from the outside environment ( ...

such as acting as a permeability barrier for most molecules and serving as the location for the transport of molecules into the cell. In addition to these functions,

prokaryotic A prokaryote () is a Unicellular organism, single-celled organism that lacks a cell nucleus, nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Greek language, Greek wikt:πρό#Ancient Greek, πρό (, 'before') a ...

membranes also function in energy conservation as the location about which a

proton motive force Chemiosmosis is the movement of ions across a semipermeable membrane bound structure, down their electrochemical gradient. An important example is the formation of adenosine triphosphate (ATP) by the movement of hydrogen ions (H+) across a membra ...

is generated. Unlike

eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...

s, bacterial membranes (with some exceptions e.g. ''

Mycoplasma ''Mycoplasma'' is a genus of bacteria that, like the other members of the class ''Mollicutes'', lack a cell wall around their cell membranes. Peptidoglycan (murein) is absent. This characteristic makes them naturally resistant to antibiotics ...

'' and methanotrophs) generally do not contain

sterol Sterol is an organic compound with formula , whose molecule is derived from that of gonane by replacement of a hydrogen atom in position 3 by a hydroxyl group. It is therefore an alcohol of gonane. More generally, any compounds that contain the go ...

s. However, many microbes do contain structurally related compounds called

hopanoid Hopanoids are a diverse subclass of triterpenoids with the same hydrocarbon skeleton as the compound hopane. This group of pentacyclic molecules therefore refers to simple hopenes, hopanols and hopanes, but also to extensively functionalized deriv ...

s which likely fulfill the same function. Unlike

eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...

s,

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 among ...

can have a wide variety of

fatty acid In chemistry, particularly in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated or unsaturated. Most naturally occurring fatty acids have an unbranched chain of an even number of carbon atoms, fr ...

s within their membranes. Along with typical saturated and unsaturated

fatty acid In chemistry, particularly in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated or unsaturated. Most naturally occurring fatty acids have an unbranched chain of an even number of carbon atoms, fr ...

s, bacteria can contain fatty acids with additional

methyl In organic chemistry, a methyl group is an alkyl derived from methane, containing one carbon atom bonded to three hydrogen atoms, having chemical formula . In formulas, the group is often abbreviated as Me. This hydrocarbon group occurs in many ...

, hydroxy or even cyclic groups. The relative proportions of these fatty acids can be modulated by the bacterium to maintain the optimum fluidity of the membrane (e.g. following temperature change). Gram-negative and mycobacteria have an inner and outer bacteria membrane. As a

phospholipid bilayer The lipid bilayer (or phospholipid bilayer) is a thin polar membrane made of two layers of lipid molecules. These membranes are flat sheets that form a continuous barrier around all cells. The cell membranes of almost all organisms and many vir ...

, the lipid portion of the

bacterial outer membrane The bacterial outer membrane is found in gram-negative bacteria. Its composition is distinct from that of the inner cytoplasmic cell membrane - among other things, the outer leaflet of the outer membrane of many gram-negative bacteria includes ...

is impermeable to charged molecules. However, channels called

porins Porins are beta barrel proteins that cross a cellular membrane and act as a pore, through which molecules can diffuse. Unlike other membrane transport proteins, porins are large enough to allow passive diffusion, i.e., they act as channels tha ...

are present in the outer membrane that allow for

passive transport Passive transport is a type of membrane transport that does not require energy to move substances across cell membranes. Instead of using cellular energy, like active transport, passive transport relies on the second law of thermodynamics to dri ...

of many

ion An ion () is an atom or molecule with a net electrical charge. The charge of an electron is considered to be negative by convention and this charge is equal and opposite to the charge of a proton, which is considered to be positive by conve ...

s,

sugar Sugar is the generic name for sweet-tasting, soluble carbohydrates, many of which are used in food. Simple sugars, also called monosaccharides, include glucose, fructose, and galactose. Compound sugars, also called disaccharides or double ...

s and

amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha am ...

s across the outer membrane. These molecules are therefore present in the

periplasm The periplasm is a concentrated gel-like matrix in the space between the inner cytoplasmic membrane and the bacterial outer membrane called the ''periplasmic space'' in gram-negative bacteria. Using cryo-electron microscopy it has been found that ...

, the region between the cytoplasmic and outer membranes. The

periplasm The periplasm is a concentrated gel-like matrix in the space between the inner cytoplasmic membrane and the bacterial outer membrane called the ''periplasmic space'' in gram-negative bacteria. Using cryo-electron microscopy it has been found that ...

contains the peptidoglycan layer and many proteins responsible for substrate binding or

hydrolysis Hydrolysis (; ) is any chemical reaction in which a molecule of water breaks one or more chemical bonds. The term is used broadly for substitution reaction, substitution, elimination reaction, elimination, and solvation reactions in which water ...

and reception of extracellular signals. The periplasm is thought to exist in a gel-like state rather than a liquid due to the high concentration of proteins and

peptidoglycan Peptidoglycan or murein is a unique large macromolecule, a polysaccharide, consisting of sugars and amino acids that forms a mesh-like peptidoglycan layer outside the plasma membrane, the rigid cell wall (murein sacculus) characteristic of most ...

found within it. Because of its location between the cytoplasmic and outer membranes, signals received and substrates bound are available to be transported across the

cytoplasmic 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 interior of all cells from the outside environment ( ...

using transport and signaling proteins imbedded there.

Extracellular (external) structures

Fimbriae and pili

Fimbriae (sometimes called " attachment pili") are protein tubes that extend out from the outer membrane in many members of the ''

Pseudomonadota Pseudomonadota (synonym Proteobacteria) is a major phylum of Gram-negative bacteria. The renaming of phyla in 2021 remains controversial among microbiologists, many of whom continue to use the earlier names of long standing in the literature. The ...

''. They are generally short in length and present in high numbers about the entire bacterial cell surface. Fimbriae usually function to facilitate the attachment of a

bacterium 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 among ...

to a surface (e.g. to form a

biofilm A biofilm comprises any syntrophic consortium of microorganisms in which cells stick to each other and often also to a surface. These adherent cells become embedded within a slimy extracellular matrix that is composed of extracellular ...

) or to other cells (e.g. animal cells during

pathogenesis Pathogenesis is the process by which a disease or disorder develops. It can include factors which contribute not only to the onset of the disease or disorder, but also to its progression and maintenance. The word comes from Greek πάθος ''pat ...

). A few organisms (e.g. ''

Myxococcus ''Myxococcus'' is a genus of bacteria in the family Myxococcaceae. Myxococci are Gram-negative, spore-forming, chemoorganotrophic, obligate aerobes. They are elongated rods with rounded or tapered ends, and they are nonflagellated. The cells mov ...

'') use fimbriae for

motility Motility is the ability of an organism to move independently, using metabolic energy. Definitions Motility, the ability of an organism to move independently, using metabolic energy, can be contrasted with sessility, the state of organisms th ...

to facilitate the assembly of multicellular structures such as

fruiting bodies The sporocarp (also known as fruiting body, fruit body or fruitbody) of fungi is a multicellular structure on which spore-producing structures, such as basidia or asci, are borne. The fruitbody is part of the sexual phase of a fungal life cyc ...

. Pili are similar in structure to fimbriae but are much longer and present on the bacterial cell in low numbers. Pili are involved in the process of

bacterial conjugation Bacterial conjugation is the transfer of genetic material between bacterial cells by direct cell-to-cell contact or by a bridge-like connection between two cells. This takes place through a pilus. It is a parasexual mode of reproduction in bacteri ...

where they are called conjugation pili or " sex pili".

Type IV pili A pilus (Latin for 'hair'; plural: ''pili'') is a hair-like appendage found on the surface of many bacteria and archaea. The terms ''pilus'' and '' fimbria'' (Latin for 'fringe'; plural: ''fimbriae'') can be used interchangeably, although some r ...

(non-sex pili) also aid bacteria in gripping surfaces.

S-layers

An

S-layer An S-layer (surface layer) is a part of the cell envelope found in almost all archaea, as well as in many types of bacteria. The S-layers of both archaea and bacteria consists of a monomolecular layer composed of only one (or, in a few cases, two) i ...

(surface layer) is a cell surface protein layer found in many different

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 among ...

and in some

archaea Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaebac ...

, where it serves as the cell wall. All

S-layer An S-layer (surface layer) is a part of the cell envelope found in almost all archaea, as well as in many types of bacteria. The S-layers of both archaea and bacteria consists of a monomolecular layer composed of only one (or, in a few cases, two) i ...

s are made up of a two-dimensional array of proteins and have a crystalline appearance, the symmetry of which differs between species. The exact function of

S-layer An S-layer (surface layer) is a part of the cell envelope found in almost all archaea, as well as in many types of bacteria. The S-layers of both archaea and bacteria consists of a monomolecular layer composed of only one (or, in a few cases, two) i ...

s is unknown, but it has been suggested that they act as a partial permeability barrier for large substrates. For example, an

S-layer An S-layer (surface layer) is a part of the cell envelope found in almost all archaea, as well as in many types of bacteria. The S-layers of both archaea and bacteria consists of a monomolecular layer composed of only one (or, in a few cases, two) i ...

could conceivably keep extracellular proteins near the cell membrane by preventing their diffusion away from the cell. In some pathogenic species, an

S-layer An S-layer (surface layer) is a part of the cell envelope found in almost all archaea, as well as in many types of bacteria. The S-layers of both archaea and bacteria consists of a monomolecular layer composed of only one (or, in a few cases, two) i ...

may help to facilitate survival within the host by conferring protection against host defence mechanisms.

Glycocalyx

Many bacteria secrete extracellular polymers outside of their cell walls called

glycocalyx The glycocalyx, also known as the pericellular matrix, is a glycoprotein and glycolipid covering that surrounds the cell membranes of bacteria, epithelial cells, and other cells. In 1970, Martinez-Palomo discovered the cell coating in animal cells ...

. These polymers are usually composed of

polysaccharide Polysaccharides (), or polycarbohydrates, are the most abundant carbohydrates found in food. They are long chain polymeric carbohydrates composed of monosaccharide units bound together by glycosidic linkages. This carbohydrate can react with wa ...

s and sometimes

protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...

. Capsules are relatively impermeable structures that cannot be stained with dyes such as India ink. They are structures that help protect bacteria from

phagocytosis Phagocytosis () is the process by which a cell uses its plasma membrane to engulf a large particle (≥ 0.5 μm), giving rise to an internal compartment called the phagosome. It is one type of endocytosis. A cell that performs phagocytosis is ...

and

desiccation Desiccation () is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its local vicinity in a moderately sealed container. ...

. Slime layer is involved in attachment of bacteria to other cells or inanimate surfaces to form

biofilm A biofilm comprises any syntrophic consortium of microorganisms in which cells stick to each other and often also to a surface. These adherent cells become embedded within a slimy extracellular matrix that is composed of extracellular ...

s. Slime layers can also be used as a food reserve for the cell.

Flagella

Perhaps the most recognizable extracellular bacterial cell structures are

flagella A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...

. Flagella are whip-like structures protruding from the bacterial cell wall and are responsible for

bacterial motility Bacterial motility is the ability of bacteria to move independently using metabolic energy. Most motility mechanisms which evolved among bacteria also evolved in parallel among the archaea. Most rod-shaped bacteria can move using their own ...

(movement). The arrangement of flagella about the bacterial cell is unique to the species observed. Common forms include: * Monotrichous – Single flagellum * Lophotrichous – A tuft of flagella found at one of the cell poles * Amphitrichous – Single flagellum found at each of two opposite poles *

Peritrichous A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...

– Multiple flagella found at several locations about the cell The bacterial flagellum consists of three basic components: a whip-like filament, a motor complex, and a hook that connects them. The filament is approximately 20 nm in diameter and consists of several protofilaments, each made up of thousands of

flagellin Flagellin is a globular protein that arranges itself in a hollow cylinder to form the filament in a bacterial flagellum. It has a mass of about 30,000 to 60,000 daltons. Flagellin is the principal component of bacterial flagella, and is present ...

subunits. The bundle is held together by a cap and may or may not be encapsulated. The motor complex consists of a series of rings anchoring the flagellum in the inner and outer membranes, followed by a proton-driven

motor An engine or motor is a machine designed to convert one or more forms of energy into mechanical energy. Available energy sources include potential energy (e.g. energy of the Earth's gravitational field as exploited in hydroelectric power g ...

that drives rotational movement in the filament.

Intracellular (internal) structures

In comparison to

eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...

s, the intracellular features of the bacterial cell are extremely simple. Bacteria do not contain

organelle In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' the ...

s in the same sense as

eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...

s. Instead, the

chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...

and perhaps

ribosome Ribosomes ( ) are macromolecular machines, found within all cells, that perform biological protein synthesis (mRNA translation). Ribosomes link amino acids together in the order specified by the codons of messenger RNA (mRNA) molecules to ...

s are the only easily observable intracellular structures found in all

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 among ...

. There do exist, however, specialized groups of bacteria that contain more complex intracellular structures, some of which are discussed below.

The bacterial DNA and plasmids

Unlike

eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...

s, the bacterial DNA is not enclosed inside of a membrane-bound

nucleus Nucleus ( : nuclei) is a Latin word for the seed inside a fruit. It most often refers to: *Atomic nucleus, the very dense central region of an atom *Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucle ...

but instead resides inside the bacterial

cytoplasm In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. The ...

. This means that the transfer of cellular information through the processes of

translation Translation is the communication of the Meaning (linguistic), meaning of a #Source and target languages, source-language text by means of an Dynamic and formal equivalence, equivalent #Source and target languages, target-language text. The ...

,

transcription Transcription refers to the process of converting sounds (voice, music etc.) into letters or musical notes, or producing a copy of something in another medium, including: Genetics * Transcription (biology), the copying of DNA into RNA, the fir ...

and

DNA replication In molecular biology, DNA replication is the biological process of producing two identical replicas of DNA from one original DNA molecule. DNA replication occurs in all living organisms acting as the most essential part for biological inheritanc ...

all occur within the same compartment and can interact with other cytoplasmic structures, most notably

ribosome Ribosomes ( ) are macromolecular machines, found within all cells, that perform biological protein synthesis (mRNA translation). Ribosomes link amino acids together in the order specified by the codons of messenger RNA (mRNA) molecules to ...

s. Bacterial DNA can be located in two places: * Bacterial chromosome, located in the irregularly shaped region known as the nucleoid * Extrachromosomal DNA, located outside of the nucleoid region as circular or linear

plasmid A plasmid is a small, extrachromosomal DNA molecule within a cell that is physically separated from chromosomal DNA and can replicate independently. They are most commonly found as small circular, double-stranded DNA molecules in bacteria; how ...

s The bacterial DNA is not packaged using

histone In biology, histones are highly basic proteins abundant in lysine and arginine residues that are found in eukaryotic cell nuclei. They act as spools around which DNA winds to create structural units called nucleosomes. Nucleosomes in turn are wr ...

s to form

chromatin Chromatin is a complex of DNA and protein found in eukaryotic cells. The primary function is to package long DNA molecules into more compact, denser structures. This prevents the strands from becoming tangled and also plays important roles in r ...

as in

eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...

s but instead exists as a highly compact

supercoil DNA supercoiling refers to the amount of twist in a particular DNA strand, which determines the amount of strain on it. A given strand may be "positively supercoiled" or "negatively supercoiled" (more or less tightly wound). The amount of a st ...

ed structure, the precise nature of which remains unclear. Most bacterial chromosomes are

circular Circular may refer to: * The shape of a circle * ''Circular'' (album), a 2006 album by Spanish singer Vega * Circular letter (disambiguation) ** Flyer (pamphlet), a form of advertisement * Circular reasoning, a type of logical fallacy * Circula ...

although some examples of linear DNA exist (e.g. ''

Borrelia burgdorferi ''Borrelia burgdorferi'' is a bacterial species of the spirochete class in the genus ''Borrelia'', and is one of the causative agents of Lyme disease in humans. Along with a few similar genospecies, some of which also cause Lyme disease, it make ...

''). Usually a single bacterial chromosome is present, although some species with multiple chromosomes have been described. Along with chromosomal DNA, most bacteria also contain small independent pieces of DNA called plasmids that often encode advantageous traits but not essential to their bacterial host. Plasmids can be easily gained or lost by a bacterium and can be transferred between bacteria as a form of

horizontal gene transfer Horizontal gene transfer (HGT) or lateral gene transfer (LGT) is the movement of genetic material between Unicellular organism, unicellular and/or multicellular organisms other than by the ("vertical") transmission of DNA from parent to offsprin ...

. So plasmids can be described as extrachromosomal DNA in a bacterial cell.

Ribosomes and other multiprotein complexes

In most

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 among ...

the most numerous intracellular structure is the

ribosome Ribosomes ( ) are macromolecular machines, found within all cells, that perform biological protein synthesis (mRNA translation). Ribosomes link amino acids together in the order specified by the codons of messenger RNA (mRNA) molecules to ...

, the site of

protein synthesis Protein biosynthesis (or protein synthesis) is a core biological process, occurring inside Cell (biology), cells, homeostasis, balancing the loss of cellular proteins (via Proteolysis, degradation or Protein targeting, export) through the product ...

in all living organisms. All

prokaryote A prokaryote () is a single-celled organism that lacks a nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Greek πρό (, 'before') and κάρυον (, 'nut' or 'kernel').Campbell, N. "Biology:Concepts & Connec ...

s have 70S (where S=

Svedberg A Svedberg unit or svedberg (symbol S, sometimes Sv) is a non- SI metric unit for sedimentation coefficients. The Svedberg unit offers a measure of a particle's size indirectly based on its sedimentation rate under acceleration (i.e. how fast a p ...

units) ribosomes while

eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...

s contain larger 80S

ribosome Ribosomes ( ) are macromolecular machines, found within all cells, that perform biological protein synthesis (mRNA translation). Ribosomes link amino acids together in the order specified by the codons of messenger RNA (mRNA) molecules to ...

s in their

cytosol The cytosol, also known as cytoplasmic matrix or groundplasm, is one of the liquids found inside cells (intracellular fluid (ICF)). It is separated into compartments by membranes. For example, the mitochondrial matrix separates the mitochondri ...

. The 70S

ribosome Ribosomes ( ) are macromolecular machines, found within all cells, that perform biological protein synthesis (mRNA translation). Ribosomes link amino acids together in the order specified by the codons of messenger RNA (mRNA) molecules to ...

is made up of a 50S and 30S subunits. The 50S subunit contains the 23S and 5S

rRNA Ribosomal ribonucleic acid (rRNA) is a type of non-coding RNA which is the primary component of ribosomes, essential to all cells. rRNA is a ribozyme which carries out protein synthesis in ribosomes. Ribosomal RNA is transcribed from ribosoma ...

while the 30S subunit contains the 16S

rRNA Ribosomal ribonucleic acid (rRNA) is a type of non-coding RNA which is the primary component of ribosomes, essential to all cells. rRNA is a ribozyme which carries out protein synthesis in ribosomes. Ribosomal RNA is transcribed from ribosoma ...

. These

rRNA Ribosomal ribonucleic acid (rRNA) is a type of non-coding RNA which is the primary component of ribosomes, essential to all cells. rRNA is a ribozyme which carries out protein synthesis in ribosomes. Ribosomal RNA is transcribed from ribosoma ...

molecules differ in size in

eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...

s and are complexed with a large number of ribosomal proteins, the number and type of which can vary slightly between organisms. While the

ribosome Ribosomes ( ) are macromolecular machines, found within all cells, that perform biological protein synthesis (mRNA translation). Ribosomes link amino acids together in the order specified by the codons of messenger RNA (mRNA) molecules to ...

is the most commonly observed intracellular multiprotein complex in

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 among ...

other large complexes do occur and can sometimes be seen using

microscopy Microscopy is the technical field of using microscopes to view objects and areas of objects that cannot be seen with the naked eye (objects that are not within the resolution range of the normal eye). There are three well-known branches of micr ...

.

Intracellular membranes

While not typical of all

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 among ...

some microbes contain intracellular membranes in addition to (or as extensions of) their cytoplasmic membranes. An early idea was that bacteria might contain membrane folds termed

mesosome Mesosomes or chondrioids are folded invaginations in the plasma membrane of bacteria that are produced by the chemical fixation techniques used to prepare samples for electron microscopy. Although several functions were proposed for these struc ...

s, but these were later shown to be artifacts produced by the chemicals used to prepare the cells for

electron microscopy An electron microscope is a microscope that uses a beam of accelerated electrons as a source of illumination. As the wavelength of an electron can be up to 100,000 times shorter than that of visible light photons, electron microscopes have a hi ...

. Examples of

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 among ...

containing intracellular membranes are

phototroph Phototrophs () are organisms that carry out photon capture to produce complex organic compounds (e.g. carbohydrates) and acquire energy. They use the energy from light to carry out various cellular metabolic processes. It is a common misconcep ...

s,

nitrifying bacteria Nitrifying bacteria are chemolithotrophic organisms that include species of genera such as ''Nitrosomonas'', ''Nitrosococcus'', ''Nitrobacter'', '' Nitrospina'', ''Nitrospira'' and '' Nitrococcus''. These bacteria get their energy from the oxidation ...

and

methane Methane ( , ) is a chemical compound with the chemical formula (one carbon atom bonded to four hydrogen atoms). It is a group-14 hydride, the simplest alkane, and the main constituent of natural gas. The relative abundance of methane on Eart ...

- oxidising bacteria. Intracellular membranes are also found in

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 among ...

belonging to the poorly studied

Planctomycetota The Planctomycetota are a phylum of widely distributed bacteria, occurring in both aquatic and terrestrial habitats. They play a considerable role in global carbon and nitrogen cycles, with many species of this phylum capable of anaerobic ammoniu ...

group, although these membranes more closely resemble organellar membranes in

eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...

s and are currently of unknown function.

Chromatophores Chromatophores are cells that produce color, of which many types are Biological pigment, pigment-containing cells, or groups of cells, found in a wide range of animals including amphibians, fish, reptiles, crustaceans and cephalopods. Mammals and ...

are intracellular membranes found in

phototroph Phototrophs () are organisms that carry out photon capture to produce complex organic compounds (e.g. carbohydrates) and acquire energy. They use the energy from light to carry out various cellular metabolic processes. It is a common misconcep ...

ic bacteria. Used primarily for photosynthesis, they contain

bacteriochlorophyll Bacteriochlorophylls (BChl) are photosynthetic pigments that occur in various phototrophic bacteria. They were discovered by C. B. van Niel in 1932. They are related to chlorophylls, which are the primary pigments in plants, algae, and cyanobact ...

pigments and carotenoids.

Cytoskeleton

The

prokaryotic cytoskeleton The prokaryotic cytoskeleton is the collective name for all structural filaments in prokaryotes. It was once thought that prokaryotic cells did not possess cytoskeletons, but advances in visualization technology and structure determination led t ...

is the collective name for all structural filaments in

prokaryote A prokaryote () is a single-celled organism that lacks a nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Greek πρό (, 'before') and κάρυον (, 'nut' or 'kernel').Campbell, N. "Biology:Concepts & Connec ...

s. It was once thought that prokaryotic cells did not possess

cytoskeleton The cytoskeleton is a complex, dynamic network of interlinking protein filaments present in the cytoplasm of all cells, including those of bacteria and archaea. In eukaryotes, it extends from the cell nucleus to the cell membrane and is compos ...

s, but advances in imaging technology and structure determination have shown the presence of filaments in these cells. Homologues for all major cytoskeletal proteins in

eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...

s have been found in prokaryotes. Cytoskeletal elements play essential roles in

cell division Cell division is the process by which a parent cell (biology), cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle in which the cell grows and replicates its chromosome(s) before dividing. In eukar ...

, protection, shape determination, and polarity determination in various prokaryotes.

Nutrient storage structures

Most

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 among ...

do not live in environments that contain large amounts of nutrients at all times. To accommodate these transient levels of nutrients

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 among ...

contain several different methods of nutrient storage in times of plenty for use in times of want. For example, many

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 among ...

store excess carbon in the form of

polyhydroxyalkanoates Polyhydroxyalkanoates or PHAs are polyesters produced in nature by numerous microorganisms, including through bacterial fermentation of sugars or lipids. When produced by bacteria they serve as both a source of energy and as a carbon store. M ...

or

glycogen Glycogen is a multibranched polysaccharide of glucose that serves as a form of energy storage in animals, fungi, and bacteria. The polysaccharide structure represents the main storage form of glucose in the body. Glycogen functions as one o ...

. Some microbes store soluble nutrients such as

nitrate Nitrate is a polyatomic ion A polyatomic ion, also known as a molecular ion, is a covalent bonded set of two or more atoms, or of a metal complex, that can be considered to behave as a single unit and that has a net charge that is not zer ...

in

vacuole A vacuole () is a membrane-bound organelle which is present in plant and fungal cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic mo ...

s. Sulfur is most often stored as elemental (S⁰) granules which can be deposited either intra- or extracellularly. Sulfur granules are especially common in

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 among ...

that use

hydrogen sulfide Hydrogen sulfide is a chemical compound with the formula . It is a colorless chalcogen-hydride gas, and is poisonous, corrosive, and flammable, with trace amounts in ambient atmosphere having a characteristic foul odor of rotten eggs. The unde ...

as an electron source. Most of the above-mentioned examples can be viewed using a

microscope A microscope () is a laboratory instrument used to examine objects that are too small to be seen by the naked eye. Microscopy is the science of investigating small objects and structures using a microscope. Microscopic means being invisibl ...

and are surrounded by a thin nonunit membrane to separate them from the

cytoplasm In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. The ...

.

Inclusions

Inclusions are considered to be nonliving components of the cell that do not possess metabolic activity and are not bounded by membranes. The most common inclusions are glycogen, lipid droplets, crystals, and pigments.

Volutin granules Volutin granules are an intracytoplasmic storage form of complexed inorganic polyphosphate, the production of which is used as one of the identifying criteria when attempting to isolate ''Corynebacterium diphtheriae'' on Löffler's medium. Polypho ...

are cytoplasmic inclusions of complexed inorganic polyphosphate. These granules are called metachromatic granules due to their displaying the metachromatic effect; they appear red or blue when stained with the blue dyes methylene blue or toluidine blue.

Gas vacuoles

Gas vacuoles Gas vesicles, also known as gas vacuoles, are nanocompartments in certain prokaryotic organisms, which help in buoyancy. Gas vesicles are composed entirely of protein; no lipids or carbohydrates have been detected. Function Gas vesicles occur ...

are membrane-bound, spindle-shaped

vesicles Vesicle may refer to: ; In cellular biology or chemistry * Vesicle (biology and chemistry), a supramolecular assembly of lipid molecules, like a cell membrane * Synaptic vesicle ; In human embryology * Vesicle (embryology), bulge-like features o ...

, found in some

plankton Plankton are the diverse collection of organisms found in Hydrosphere, water (or atmosphere, air) that are unable to propel themselves against a Ocean current, current (or wind). The individual organisms constituting plankton are called plankt ...

ic bacteria and ''

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, blu ...

'', that provides

buoyancy Buoyancy (), or upthrust, is an upward force exerted by a fluid that opposes the weight of a partially or fully immersed object. In a column of fluid, pressure increases with depth as a result of the weight of the overlying fluid. Thus the p ...

to these cells by decreasing their overall cell

density Density (volumetric mass density or specific mass) is the substance's mass per unit of volume. The symbol most often used for density is ''ρ'' (the lower case Greek letter rho), although the Latin letter ''D'' can also be used. Mathematical ...

. Positive buoyancy is needed to keep the cells in the upper reaches of the water column, so that they can continue to perform

photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored i ...

. They are made up of a shell of protein that has a highly

hydrophobic In chemistry, hydrophobicity is the physical property of a molecule that is seemingly repelled from a mass of water (known as a hydrophobe). In contrast, hydrophiles are attracted to water. Hydrophobic molecules tend to be nonpolar and, th ...

inner surface, making it impermeable to water (and stopping water vapour from condensing inside) but permeable to most

gas Gas is one of the four fundamental states of matter (the others being solid, liquid, and plasma). A pure gas may be made up of individual atoms (e.g. a noble gas like neon), elemental molecules made from one type of atom (e.g. oxygen), or ...

es. Because the gas vesicle is a hollow cylinder, it is liable to collapse when the surrounding

pressure Pressure (symbol: ''p'' or ''P'') is the force applied perpendicular to the surface of an object per unit area over which that force is distributed. Gauge pressure (also spelled ''gage'' pressure)The preferred spelling varies by country and e ...

increases. Natural selection has fine tuned the structure of the gas vesicle to maximise its resistance to

buckling In structural engineering, buckling is the sudden change in shape (deformation) of a structural component under load, such as the bowing of a column under compression or the wrinkling of a plate under shear. If a structure is subjected to a gr ...

, including an external strengthening protein, GvpC, rather like the green thread in a braided hosepipe. There is a simple relationship between the diameter of the gas vesicle and pressure at which it will collapse – the wider the gas vesicle the weaker it becomes. However, wider gas vesicles are more efficient, providing more buoyancy per unit of protein than narrow gas vesicles. Different species produce gas vesicle of different diameter, allowing them to colonise different depths of the water column (fast growing, highly competitive species with wide gas vesicles in the top most layers; slow growing, dark-adapted, species with strong narrow gas vesicles in the deeper layers). The diameter of the gas vesicle will also help determine which species survive in different bodies of water. Deep lakes that experience winter mixing expose the cells to the hydrostatic pressure generated by the full water column. This will select for species with narrower, stronger gas vesicles. The cell achieves its height in the water column by synthesising gas vesicles. As the cell rises up, it is able to increase its

carbohydrate In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or ma ...

load through increased photosynthesis. Too high and the cell will suffer photobleaching and possible death, however, the carbohydrate produced during photosynthesis increases the cell's density, causing it to sink. The daily cycle of carbohydrate build-up from photosynthesis and carbohydrate

catabolism Catabolism () is the set of metabolic pathways that breaks down molecules into smaller units that are either oxidized to release energy or used in other anabolic reactions. Catabolism breaks down large molecules (such as polysaccharides, lipids, ...

during dark hours is enough to fine-tune the cell's position in the water column, bring it up toward the surface when its carbohydrate levels are low and it needs to photosynthesis, and allowing it to sink away from the harmful

UV radiation Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30 PHz) to 400 nm (750 THz), shorter than that of visible light, but longer than X-rays. UV radiation i ...

when the cell's carbohydrate levels have been replenished. An extreme excess of carbohydrate causes a significant change in the internal pressure of the cell, which causes the gas vesicles to buckle and collapse and the cell to sink out.

Microcompartments

Bacterial microcompartment Bacterial microcompartments (BMCs) are organelle-like structures found in bacteria. They consist of a protein shell that encloses enzymes and other proteins. BMCs are typically about 40–200 nanometers in diameter and are made entirely of protei ...

s are widespread, membrane-bound organelles that are made of a protein shell that surrounds and encloses various enzymes. provide a further level of organization; they are compartments within bacteria that are surrounded by polyhedral protein shells, rather than by lipid membranes. These "polyhedral organelles" localize and compartmentalize bacterial metabolism, a function performed by the membrane-bound organelles in eukaryotes.

= Carboxysomes

=

Carboxysome Carboxysomes are bacterial microcompartments (BMCs) consisting of polyhedral protein shells filled with the enzymes ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBisCO)—the predominant enzyme in carbon fixation and the rate limiting e ...

s are bacterial microcompartments found in many

autotrophic An autotroph or primary producer is an organism that produces complex organic compounds (such as carbohydrates, fats, and proteins) using carbon from simple substances such as carbon dioxide,Morris, J. et al. (2019). "Biology: How Life Works", ...

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 among ...

such as Cyanobacteria, Knallgasbacteria, Nitroso- and Nitrobacteria. They are proteinaceous structures resembling phage heads in their

morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines * Morphology (archaeology), study of the shapes or forms of artifacts * Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies ...

and contain the enzymes of carbon dioxide fixation in these organisms (especially ribulose bisphosphate carboxylase/oxygenase, RuBisCO, and carbonic anhydrase). It is thought that the high local concentration of the enzymes along with the fast conversion of bicarbonate to carbon dioxide by carbonic anhydrase allows faster and more efficient carbon dioxide fixation than possible inside the cytoplasm. Similar structures are known to harbor the coenzyme B12-containing

glycerol dehydratase In enzymology, a glycerol dehydratase () catalyzes the chemical reaction :glycerol \rightleftharpoons 3-hydroxypropanal + H2O This enzyme belongs to the family of lyases, specifically the hydro-lyases, which cleave carbon-oxygen bonds. The s ...

, the key enzyme of glycerol fermentation to 1,3-propanediol, in some Enterobacteriaceae (e. g. Salmonella).

= Magnetosomes

=

Magnetosome Magnetosomes are membranous structures present in magnetotactic bacteria (MTB). They contain iron-rich magnetic particles that are enclosed within a lipid bilayer membrane. Each magnetosome can often contain 15 to 20 magnetite crystals that form a ...

s are bacterial microcompartments found in

magnetotactic bacteria Magnetotactic bacteria (or MTB) are a polyphyletic group of bacteria that orient themselves along the magnetic field lines of Earth's magnetic field. Discovered in 1963 by Salvatore Bellini and rediscovered in 1975 by Richard Blakemore, this alig ...

that allow them to sense and align themselves along a magnetic field (

magnetotaxis Magnetotaxis is a process implemented by a diverse group of Gram-negative bacteria that involves orienting and coordinating movement in response to Earth's magnetic field. This process is mainly carried out by microaerophilic and anaerobic bacteri ...

). The ecological role of magnetotaxis is unknown but is thought to be involved in the determination of optimal oxygen concentrations. Magnetosomes are composed of the mineral

magnetite Magnetite is a mineral and one of the main iron ores, with the chemical formula Fe2+Fe3+2O4. It is one of the oxides of iron, and is ferrimagnetic; it is attracted to a magnet and can be magnetized to become a permanent magnet itself. With the ...

or

greigite Greigite is an iron sulfide mineral with the chemical formula . It is the sulfur equivalent of the iron oxide magnetite (Fe3O4). It was first described in 1964 for an occurrence in San Bernardino County, California, and named after the mineralogis ...

and are surrounded by a lipid bilayer membrane. The morphology of magnetosomes is species-specific.

Endospores

Perhaps the best known bacterial adaptation to stress is the formation of

endospore An endospore is a dormant, tough, and non-reproductive structure produced by some bacteria in the phylum Bacillota. The name "endospore" is suggestive of a spore or seed-like form (''endo'' means 'within'), but it is not a true spore (i.e., no ...

s. Endospores are bacterial survival structures that are highly resistant to many different types of chemical and environmental stresses and therefore enable the survival of

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 among ...

in environments that would be lethal for these cells in their normal vegetative form. It has been proposed that endospore formation has allowed for the survival of some bacteria for hundreds of millions of years (e.g. in salt crystals) although these publications have been questioned. Endospore formation is limited to several genera of gram-positive bacteria such as ''

Bacillus ''Bacillus'' (Latin "stick") is a genus of Gram-positive, rod-shaped bacteria, a member of the phylum ''Bacillota'', with 266 named species. The term is also used to describe the shape (rod) of other so-shaped bacteria; and the plural ''Bacilli ...

'' and '' Clostridium''. It differs from reproductive spores in that only one spore is formed per cell resulting in no net gain in cell number upon endospore germination. The location of an endospore within a cell is species-specific and can be used to determine the identity of a bacterium.

Dipicolinic acid Dipicolinic acid (pyridine-2,6-dicarboxylic acid or PDC and DPA) is a chemical compound which plays a role in the heat resistance of bacterial endospores. It is also used to prepare dipicolinato ligated lanthanide and transition metal complexes f ...

is a chemical compound which composes 5% to 15% of the dry weight of bacterial spores and is implicated in being responsible for the heat resistance of endospores. Archaeologists have found viable endospores taken from the intestines of Egyptian mummies as well as from lake sediments in Northern Sweden estimated to be many thousands of years old.

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{{Anatomy Bacteria Bacteriology

Cell morphology

Cell wall

The gram-positive cell wall

The gram-negative cell wall

Plasma membrane

Extracellular (external) structures

Fimbriae and pili

S-layers

Glycocalyx

Flagella

Intracellular (internal) structures

The bacterial DNA and plasmids

Ribosomes and other multiprotein complexes

Intracellular membranes

Cytoskeleton

Nutrient storage structures

Inclusions

Gas vacuoles

Microcompartments

= Carboxysomes

= Magnetosomes

Endospores

References

Further reading

External links