Cell
Onion (Allium cepa) root cells in different phases of the cell cycle (drawn by E. B. Wilson, 1900)
A eukaryotic cell (left) and prokaryotic cell (right)
Identifiers
MeSHD002477
THH1.00.01.0.00001
FMA686465
Anatomical terminology

The cell is the basic structural and functional unit of all forms of life. Every cell consists of cytoplasm enclosed within a membrane, and contains many macromolecules such as proteins, DNA and RNA, as well as many small molecules of nutrients and metabolites.[1] The term comes from the Latin word cellula meaning 'small room'.[2]

Cells can acquire specified function and carry out various tasks within the cell such as replication, DNA repair, protein synthesis, and motility. Cells are capable of specialization and mobility within the cell.

Most plant and animal cells are only visible under a light microscope, with dimensions between 1 and 100 micrometres.[3] Electron microscopy gives a much higher resolution showing greatly detailed cell structure. Organisms can be classified as unicellular (consisting of a single cell such as bacteria) or multicellular (including plants and animals).[4] Most unicellular organisms are classed as microorganisms. The number of cells in plants and animals varies from species to species; it has been estimated that the human body contains around 37 trillion (3.72×1013) cells.[5] The human brain accounts for around 80 billion of these cells.[6]

The study of cells and how they work has led to many other studies in related areas of biology, including: discovery of DNA, cancer systems biology, aging and developmental biology.

Cell biology is the study of cells, which were discovered by Robert Hooke in 1665, who named them for their resemblance to cells inhabited by Christian monks in a monastery.[7][8] Cell theory, first developed in 1839 by Matthias Jakob Schleiden and Theodor Schwann, states that all organisms are composed of one or more cells, that cells are the fundamental unit of structure and function in all living organisms, and that all cells come from pre-existing cells.[9] Cells emerged on Earth about 4 billion years ago.[10][11][12][13]

Discovery

Main article: Cell theory

With continual improvements made to microscopes over time, magnification technology became advanced enough to discover cells. This discovery is largely attributed to Robert Hooke, and began the scientific study of cells, known as cell biology. When observing a piece of cork under the scope, he was able to see pores. This was shocking at the time as it was believed no one else had seen these. To further support his theory, Matthias Schleiden and Theodor Schwann both also studied cells of both animal and plants. What they discovered were significant differences between the two types of cells. This put forth the idea that cells were not only fundamental to plants, but animals as well.

Cell types

Cells are broadly categorized into two types: eukaryotic cells, which possesses a nucleus, and prokaryotic cells, which lack a nucleus but still has a nucleoid region. Prokaryotes are single-celled organisms, whereas eukaryotes can be either single-celled or multicellular.[14]

Prokaryotic cells

Main article: Prokaryote

Structure of a typical prokaryotic cell

Prokaryotes include bacteria and archaea, two of the three domains of life. Prokaryotic cells were the first form of life on Earth, characterized by having vital biological processes including cell signaling. They are simpler and smaller than eukaryotic cells, and lack a nucleus, and other membrane-bound organelles. The DNA of a prokaryotic cell consists of a single circular chromosome that is in direct contact with the cytoplasm. The nuclear region in the cytoplasm is called the nucleoid. Most prokaryotes are the smallest of all organisms ranging from 0.5 to 2.0 μm in diameter.[15]

A prokaryotic cell has three regions:

Bacterial shapes

Main article: Bacteria § Morphology

Cell shape, also called cell morphology, has been hypothesized to form from the arrangement and movement of the cytoskeleton.[17] Many advancements in the study of cell morphology come from studying simple bacteria such as Staphylococcus aureusE. coli, and B. subtilis.[18] Different cell shapes have been found and described, but how and why cells form different shapes is still widely unknown.[18] Some cell shapes that have been identified include rods, cocci and spirochaetes. Cocci have a circular shape, bacilli have an elongated rod-like shape, and spirochaetes have a spiral shape. Many other shapes have also been determined.

Eukaryotic cells

Main article: Eukaryote

Structure of a typical animal cell
Structure of a typical plant cell

Plants, animals, fungi, slime moulds, protozoa, and algae are all eukaryotic. These cells are about fifteen times wider than a typical prokaryote and can be as much as a thousand times greater in volume. The main distinguishing feature of eukaryotes as compared to prokaryotes is compartmentalization: the presence of membrane-bound organelles (compartments) in which specific activities take place. Most important among these is a cell nucleus,[4] an organelle that houses the cell's DNA. This nucleus gives the eukaryote its name, which means "true kernel (nucleus)". Some of the other differences are:

Comparison of features of prokaryotic and eukaryotic cells
Prokaryotes Eukaryotes
Typical organisms bacteria, archaea protists, fungi, plants, animals
Typical size ~ 1–5 μm[22] ~ 10–100 μm[22]
Type of nucleus nucleoid region; no true nucleus true nucleus with double membrane
DNA circular (usually) linear molecules (chromosomes) with histone proteins
RNA/protein synthesis coupled in the cytoplasm RNA synthesis in the nucleus
protein synthesis in the cytoplasm
Ribosomes 50S and 30S 60S and 40S
Cytoplasmic structure very few structures highly structured by endomembranes and a cytoskeleton
Cell movement flagella made of flagellin flagella and cilia containing microtubules; lamellipodia and filopodia containing actin
Mitochondria none one to several thousand
Chloroplasts none in algae and plants
Organization usually single cells single cells, colonies, higher multicellular organisms with specialized cells
Cell division binary fission (simple division) mitosis (fission or budding)
meiosis
Chromosomes single chromosome more than one chromosome
Membranes cell membrane

Subcellular components

All cells, whether prokaryotic or eukaryotic, have a membrane that envelops the cell, regulates what moves in and out (selectively permeable), and maintains the electric potential of the cell. Inside the membrane, the cytoplasm takes up most of the cell's volume. Except red blood cells, which lack a cell nucleus and most organelles to accommodate maximum space for hemoglobin, all cells possess DNA, the hereditary material of genes, and RNA, containing the information necessary to build various proteins such as enzymes, the cell's primary machinery. There are also other kinds of biomolecules in cells. This article lists these primary cellular components, then briefly describes their function.

Cell membrane

Main article: Cell membrane

Detailed diagram of lipid bilayer of cell membrane

The cell membrane, or plasma membrane, is a selectively permeable[23] biological membrane that surrounds the cytoplasm of a cell. In animals, the plasma membrane is the outer boundary of the cell, while in plants and prokaryotes it is usually covered by a cell wall. This membrane serves to separate and protect a cell from its surrounding environment and is made mostly from a double layer of phospholipids, which are amphiphilic (partly hydrophobic and partly hydrophilic). Hence, the layer is called a phospholipid bilayer, or sometimes a fluid mosaic membrane. Embedded within this membrane is a macromolecular structure called the porosome the universal secretory portal in cells and a variety of protein molecules that act as channels and pumps that move different molecules into and out of the cell.[4] The membrane is semi-permeable, and selectively permeable, in that it can either let a substance (molecule or ion) pass through freely, pass through to a limited extent or not pass through at all. Cell surface membranes also contain receptor proteins that allow cells to detect external signaling molecules such as hormones.

Cytoskeleton

Main article: Cytoskeleton

Further information: Morphogenesis

A fluorescent image of an endothelial cell. Nuclei are stained blue, mitochondria are stained red, and microfilaments are stained green.

The cytoskeleton acts to organize and maintain the cell's shape; anchors organelles in place; helps during endocytosis, the uptake of external materials by a cell, and cytokinesis, the separation of daughter cells after cell division; and moves parts of the cell in processes of growth and mobility. The eukaryotic cytoskeleton is composed of microtubules, intermediate filaments and microfilaments. In the cytoskeleton of a neuron the intermediate filaments are known as neurofilaments. There are a great number of proteins associated with them, each controlling a cell's structure by directing, bundling, and aligning filaments.[4] The prokaryotic cytoskeleton is less well-studied but is involved in the maintenance of cell shape, polarity and cytokinesis.[24] The subunit protein of microfilaments is a small, monomeric protein called actin. The subunit of microtubules is a dimeric molecule called tubulin. Intermediate filaments are heteropolymers whose subunits vary among the cell types in different tissues. Some of the subunit proteins of intermediate filaments include vimentin, desmin, lamin (lamins A, B and C), keratin (multiple acidic and basic keratins), and neurofilament proteins (NF–L, NF–M).

Genetic material

Main articles: DNA and RNA

Deoxyribonucleic acid (DNA)

Two different kinds of genetic material exist: deoxyribonucleic acid (DNA) and ribonucleic acid (RNA). Cells use DNA for their long-term information storage. The biological information contained in an organism is encoded in its DNA sequence.[4] RNA is used for information transport (e.g., mRNA) and enzymatic functions (e.g., ribosomal RNA). Transfer RNA (tRNA) molecules are used to add amino acids during protein translation.

Prokaryotic genetic material is organized in a simple circular bacterial chromosome in the nucleoid region of the cytoplasm. Eukaryotic genetic material is divided into different,[4] linear molecules called chromosomes inside a discrete nucleus, usually with additional genetic material in some organelles like mitochondria and chloroplasts (see endosymbiotic theory).

A human cell has genetic material contained in the cell nucleus (the nuclear genome) and in the mitochondria (the mitochondrial genome). In humans, the nuclear genome is divided into 46 linear DNA molecules called chromosomes, including 22 homologous chromosome pairs and a pair of sex chromosomes. The mitochondrial genome is a circular DNA molecule distinct from nuclear DNA. Although the mitochondrial DNA is very small compared to nuclear chromosomes,[4] it codes for 13 proteins involved in mitochondrial energy production and specific tRNAs.

Foreign genetic material (most commonly DNA) can also be artificially introduced into the cell by a process called transfection. This can be transient, if the DNA is not inserted into the cell's genome, or stable, if it is. Certain viruses also insert their genetic material into the genome.

Organelles

Main article: Organelle

Organelles are parts of the cell that are adapted and/or specialized for carrying out one or more vital functions, analogous to the organs of the human body (such as the heart, lung, and kidney, with each organ performing a different function).[4] Both eukaryotic and prokaryotic cells have organelles, but prokaryotic organelles are generally simpler and are not membrane-bound.

There are several types of organelles in a cell. Some (such as the nucleus and Golgi apparatus) are typically solitary, while others (such as mitochondria, chloroplasts, peroxisomes and lysosomes) can be numerous (hundreds to thousands). The cytosol is the gelatinous fluid that fills the cell and surrounds the organelles.

Eukaryotic

Human cancer cells, specifically HeLa cells, with DNA stained blue. The central and rightmost cell are in interphase, so their DNA is diffuse and the entire nuclei are labelled. The cell on the left is going through mitosis and its chromosomes have condensed.
Diagram of the endomembrane system

Eukaryotic and prokaryotic

Structures outside the cell membrane

Many cells also have structures which exist wholly or partially outside the cell membrane. These structures are notable because they are not protected from the external environment by the semipermeable cell membrane. In order to assemble these structures, their components must be carried across the cell membrane by export processes.

Cell wall

Further information: Cell wall

Many types of prokaryotic and eukaryotic cells have a cell wall. The cell wall acts to protect the cell mechanically and chemically from its environment, and is an additional layer of protection to the cell membrane. Different types of cell have cell walls made up of different materials; plant cell walls are primarily made up of cellulose, fungi cell walls are made up of chitin and bacteria cell walls are made up of peptidoglycan.

Prokaryotic

Capsule

A gelatinous capsule is present in some bacteria outside the cell membrane and cell wall. The capsule may be polysaccharide as in pneumococci, meningococci or polypeptide as Bacillus anthracis or hyaluronic acid as in streptococci. Capsules are not marked by normal staining protocols and can be detected by India ink or methyl blue, which allows for higher contrast between the cells for observation.[26]: 87 

Flagella

Flagella are organelles for cellular mobility. The bacterial flagellum stretches from cytoplasm through the cell membrane(s) and extrudes through the cell wall. They are long and thick thread-like appendages, protein in nature. A different type of flagellum is found in archaea and a different type is found in eukaryotes.

Fimbriae

A fimbria (plural fimbriae also known as a pilus, plural pili) is a short, thin, hair-like filament found on the surface of bacteria. Fimbriae are formed of a protein called pilin (antigenic) and are responsible for the attachment of bacteria to specific receptors on human cells (cell adhesion). There are special types of pili involved in bacterial conjugation.

Cellular processes

Prokaryotes divide by binary fission, while eukaryotes divide by mitosis or meiosis.

Replication

Main article: Cell division

Cell division involves a single cell (called a mother cell) dividing into two daughter cells. This leads to growth in multicellular organisms (the growth of tissue) and to procreation (vegetative reproduction) in unicellular organisms. Prokaryotic cells divide by binary fission, while eukaryotic cells usually undergo a process of nuclear division, called mitosis, followed by division of the cell, called cytokinesis. A diploid cell may also undergo meiosis to produce haploid cells, usually four. Haploid cells serve as gametes in multicellular organisms, fusing to form new diploid cells.

DNA replication, or the process of duplicating a cell's genome,[4] always happens when a cell divides through mitosis or binary fission. This occurs during the S phase of the cell cycle.

In meiosis, the DNA is replicated only once, while the cell divides twice. DNA replication only occurs before meiosis I. DNA replication does not occur when the cells divide the second time, in meiosis II.[27] Replication, like all cellular activities, requires specialized proteins for carrying out the job.[4]

An outline of the catabolism of proteins, carbohydrates and fats

DNA repair

Main article: DNA repair

In general, cells of all organisms contain enzyme systems that scan their DNA for DNA damage and carry out repair processes when damage is detected.[28] Diverse repair processes have evolved in organisms ranging from bacteria to humans. The widespread prevalence of these repair processes indicates the importance of maintaining cellular DNA in an undamaged state in order to avoid cell death or errors of replication due to damage that could lead to mutation. E. coli bacteria are a well-studied example of a cellular organism with diverse well-defined DNA repair processes. These include: (1) nucleotide excision repair, (2) DNA mismatch repair, (3) non-homologous end joining of double-strand breaks, (4) recombinational repair and (5) light-dependent repair (photoreactivation).

Growth and metabolism

An overview of protein synthesis.
Within the nucleus of the cell (light blue), genes (DNA, dark blue) are transcribed into RNA. This RNA is then subject to post-transcriptional modification and control, resulting in a mature mRNA (red) that is then transported out of the nucleus and into the cytoplasm (peach), where it undergoes translation into a protein. mRNA is translated by ribosomes (purple) that match the three-base codons of the mRNA to the three-base anti-codons of the appropriate tRNA. Newly synthesized proteins (black) are often further modified, such as by binding to an effector molecule (orange), to become fully active.

Main articles: Cell growth and Metabolism

Between successive cell divisions, cells grow through the functioning of cellular metabolism. Cell metabolism is the process by which individual cells process nutrient molecules. Metabolism has two distinct divisions: catabolism, in which the cell breaks down complex molecules to produce energy and reducing power, and anabolism, in which the cell uses energy and reducing power to construct complex molecules and perform other biological functions. Complex sugars consumed by the organism can be broken down into simpler sugar molecules called monosaccharides such as glucose. Once inside the cell, glucose is broken down to make adenosine triphosphate (ATP),[4] a molecule that possesses readily available energy, through two different pathways.

Protein synthesis

Main article: Protein biosynthesis

Cells are capable of synthesizing new proteins, which are essential for the modulation and maintenance of cellular activities. This process involves the formation of new protein molecules from amino acid building blocks based on information encoded in DNA/RNA. Protein synthesis generally consists of two major steps: transcription and translation.

Transcription is the process where genetic information in DNA is used to produce a complementary RNA strand. This RNA strand is then processed to give messenger RNA (mRNA), which is free to migrate through the cell. mRNA molecules bind to protein-RNA complexes called ribosomes located in the cytosol, where they are translated into polypeptide sequences. The ribosome mediates the formation of a polypeptide sequence based on the mRNA sequence. The mRNA sequence directly relates to the polypeptide sequence by binding to transfer RNA (tRNA) adapter molecules in binding pockets within the ribosome. The new polypeptide then folds into a functional three-dimensional protein molecule.

Motility

Main article: Motility

Unicellular organisms can move in order to find food or escape predators. Common mechanisms of motion include flagella and cilia.

In multicellular organisms, cells can move during processes such as wound healing, the immune response and cancer metastasis. For example, in wound healing in animals, white blood cells move to the wound site to kill the microorganisms that cause infection. Cell motility involves many receptors, crosslinking, bundling, binding, adhesion, motor and other proteins.[29] The process is divided into three steps: protrusion of the leading edge of the cell, adhesion of the leading edge and de-adhesion at the cell body and rear, and cytoskeletal contraction to pull the cell forward. Each step is driven by physical forces generated by unique segments of the cytoskeleton.[30][31]

Navigation, control and communication

See also: Cybernetics § In biology

In August 2020, scientists described one way cells—in particular cells of a slime mold and mouse pancreatic cancer-derived cells—are able to navigate efficiently through a body and identify the best routes through complex mazes: generating gradients after breaking down diffused chemoattractants which enable them to sense upcoming maze junctions before reaching them, including around corners.[32][33][34]

Multicellularity

Main article: Multicellular organism

Cell specialization/differentiation

Main article: Cellular differentiation

Staining of a Caenorhabditis elegans highlights the nuclei of its cells.

Multicellular organisms are organisms that consist of more than one cell, in contrast to single-celled organisms.[35]

In complex multicellular organisms, cells specialize into different cell types that are adapted to particular functions. In mammals, major cell types include skin cells, muscle cells, neurons, blood cells, fibroblasts, stem cells, and others. Cell types differ both in appearance and function, yet are genetically identical. Cells are able to be of the same genotype but of different cell type due to the differential expression of the genes they contain.

Most distinct cell types arise from a single totipotent cell, called a zygote, that differentiates into hundreds of different cell types during the course of development. Differentiation of cells is driven by different environmental cues (such as cell–cell interaction) and intrinsic differences (such as those caused by the uneven distribution of molecules during division).

Origin of multicellularity

Multicellularity has evolved independently at least 25 times,[36] including in some prokaryotes, like cyanobacteria, myxobacteria, actinomycetes, Magnetoglobus multicellularis, or Methanosarcina. However, complex multicellular organisms evolved only in six eukaryotic groups: animals, fungi, brown algae, red algae, green algae, and plants.[37] It evolved repeatedly for plants (Chloroplastida), once or twice for animals, once for brown algae, and perhaps several times for fungi, slime molds, and red algae.[38] Multicellularity may have evolved from colonies of interdependent organisms, from cellularization, or from organisms in symbiotic relationships.

The first evidence of multicellularity is from cyanobacteria-like organisms that lived between 3 and 3.5 billion years ago.[36] Other early fossils of multicellular organisms include the contested Grypania spiralis and the fossils of the black shales of the Palaeoproterozoic Francevillian Group Fossil B Formation in Gabon.[39]

The evolution of multicellularity from unicellular ancestors has been replicated in the laboratory, in evolution experiments using predation as the selective pressure.[36]

Origins

Main article: Evolutionary history of life

The origin of cells has to do with the origin of life, which began the history of life on Earth.

Origin of the first cell

Stromatolites are left behind by cyanobacteria, also called blue-green algae. They are the oldest known fossils of life on Earth. This one-billion-year-old fossil is from Glacier National Park in the United States.

Further information: Abiogenesis and Evolution of cells

There are several theories about the origin of small molecules that led to life on the early Earth. They may have been carried to Earth on meteorites (see Murchison meteorite), created at deep-sea vents, or synthesized by lightning in a reducing atmosphere (see Miller–Urey experiment). There is little experimental data defining what the first self-replicating forms were. RNA is thought to be the earliest self-replicating molecule, as it is capable of both storing genetic information and catalyzing chemical reactions (see RNA world hypothesis), but some other entity with the potential to self-replicate could have preceded RNA, such as clay or peptide nucleic acid.[40]

Cells emerged at least 3.5 billion years ago.[41][42][43] The current belief is that these cells were heterotrophs. The early cell membranes were probably more simple and permeable than modern ones, with only a single fatty acid chain per lipid. Lipids are known to spontaneously form bilayered vesicles in water, and could have preceded RNA, but the first cell membranes could also have been produced by catalytic RNA, or even have required structural proteins before they could form.[44]

Origin of eukaryotic cells

Further information: Evolution of sexual reproduction

The eukaryotic cell seems to have evolved from a symbiotic community of prokaryotic cells. DNA-bearing organelles like the mitochondria and the chloroplasts are descended from ancient symbiotic oxygen-breathing Alphaproteobacteria and "Cyanobacteria", respectively, which were endosymbiosed by an ancestral archaean prokaryote.

There is still considerable debate about whether organelles like the hydrogenosome predated the origin of mitochondria, or vice versa: see the hydrogen hypothesis for the origin of eukaryotic cells.

History of research

Main article: Cell theory

Robert Hooke's drawing of cells in cork, 1665

See also

References

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