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Cell

The cell — the basic structural and functional unit of every living organism: its discovery under the microscope, its organelles, the split between prokaryotes and eukaryotes, and how cells grow and divide.

The cell is the smallest unit of life, and all organisms are built from one or more of them. Robert Hooke coined the term in 1665 after observing cork through a microscope, and in 1838 and 1839 Matthias Schleiden and Theodor Schwann proposed that plants and animals alike are made of cells, a theory completed by Rudolf Virchow's principle that all cells arise from existing cells. Cells come in two great classes — prokaryotes, which lack a nuclear envelope, and eukaryotes, whose DNA is enclosed in a nucleus alongside membrane-bound organelles such as mitochondria and chloroplasts, both descended from symbiotic bacteria. Cells reproduce by division: the mitotic cell cycle passes through G1, S, G2 and M phases, while meiosis halves the chromosome number to produce gametes.

Dozens of elongated cultured cells glowing cyan-green on a black background, with red filament fringes and violet-blue oval nuclei.
Multiphoton fluorescence image of cultured HeLa cells, with actin filaments in red, microtubules in cyan and cell nuclei in blue (NIH). · National Institutes of Health (NIH) · Public domain
Red blood cell diameterAbout eight micrometers (8 μm)
Bacterial cell sizeMost bacterial cells are 1 to 10 μm in diameter; spherical, rod-shaped or spiral
In this article
  1. 1Overview
  2. 2Cell Theory
  3. 3Historical Observation of Cells
  4. 4Microscopy and the Study of Cells
  5. 5Universal Features and the Prokaryote–Eukaryote Divide
  6. 6Eukaryotic Cell Structure
  7. 7Cell Size and Scale
  8. 8Genomes and Ribosome Counts
  9. 9Origins and Evolution
  10. 10The Cell Cycle
  11. 11Control of the Cell Cycle
  12. 12Cell Division, Meiosis and Sexual Reproduction
  13. 13Timeline: from Hooke's Microscope to Super-Resolution Microscopy
  14. 14Frequently asked questions
  15. 15Knowledge graph

Overview

A cell is, in biology, the basic membrane-bound unit that contains the fundamental molecules of life and of which all living things are composed. It is enclosed by a plasma membrane, a selective barrier that allows nutrients to enter and waste products to leave. Cells are the basic building blocks of all organisms, and the human body is composed of trillions of cells.

1. Cell Theory

Cell theory is the fundamental scientific theory of biology holding that cells are the basic units of all living tissues. Its modern formulation rests on two tenets: biogenesis, the principle that all cells come only from other cells, and the recognition of cells as the fundamental units of organisms. The cell doctrine states that cells are generated from cells, and that the only way to make more cells is by division of those that already exist.

The theory was first proposed by Theodor Schwann and Matthias Jakob Schleiden in 1838. Schwann's microscopic investigations showed that animal tissues, like those of plants, are composed of cells, and that each cell has a nucleus. Schleiden, by contrast, believed that cells formed through crystallization rather than cell division. With the atlas texts of histopathology published in the 1840s, the cell theory introduced and publicized by Schwann in 1838 and 1839 became universally accepted as scientific fact.

In 1852 Robert Remak published convincing evidence that cells are derived from other cells as a result of cell division. Rudolf Virchow published an editorial essay entitled Cellular Pathology, which popularized the concept of cell theory using the Latin phrase omnis cellula a cellula, meaning that all cells arise from cells. Remak, a former colleague who worked in the same laboratory as Virchow at the University of Berlin, had published the same idea three years before Virchow's essay. Virchow was one of the first to determine the causes of various diseases by examining their effects on tissues and organs, and over the course of his career he published more than 2,000 papers.

2. Historical Observation of Cells

  • 1665
    Hooke describes cork and names the cell
    Robert Hooke described cork and other plant tissues in 1665 and introduced the term cell because the cellulose walls of dead cork cells reminded him of the blocks of cells occupied by monks.4Cell theory | Definition, History, Importance, Scientists, First Proposed, & Facts · Encyclopaedia Britannica ↗
  • 1831
    Brown first describes nuclei
    The Scottish botanist Robert Brown was the first to describe observations of nuclei, which he observed in plant cells.7Microbiology, 3.2 Foundations of Modern Cell Theory · OpenStax, Rice University ↗
  • 1833
    The nucleus recognized as a constant component
    The nucleus was observed by the Scottish botanist Robert Brown in 1833 as a constant component of plant cells.4
Seventeenth-century black-and-white engraving: a dark circle packed with honeycomb-like walled pores, and a small cork twig below.
Robert Hooke's engraving of cork from Micrographia (1667): the honeycomb-like pores he named 'cells', shown with a cork branch at the foot of the plate. · National Library of Medicine · Public domain

3. Microscopy and the Study of Cells

The reach of any microscope is bounded by two distinct limits: how much it can enlarge an object and how fine a detail it can separate. A simple microscope can resolve below 1 micrometre, while a compound microscope can resolve down to about 0.2 μm. In terms of enlargement, single-lensed simple microscopes can magnify up to 300 times and are capable of revealing bacteria, whereas compound microscopes can magnify up to 2,000 times.

Antonie van Leeuwenhoek learned to grind lenses before 1668 and used them to make simple hand-held microscopes that, with appropriate lighting, allowed him to magnify objects over 275 times. His instruments employed lenses ranging from 70x to over 250x.

The transmission electron microscope has magnifying powers of more than 1,000,000 times and forms images of thin specimens, typically sections, in a near vacuum. The instrument was first developed by Ernst Ruska in 1933.

4. Universal Features and the Prokaryote–Eukaryote Divide

The chief component of prokaryotic cell walls is peptidoglycan, while the major organic molecule in the plant cell wall is cellulose, a polysaccharide comprised of glucose units. Many bacteria are further enclosed by a polysaccharide capsule, and some prokaryotes bear flagella, pili or fimbriae. These envelopes and appendages distinguish the prokaryotic surface from the cellulose-based wall of the plant cell.

Brown-pink rod-shaped bacteria heaped in a cluster on a rough surface, with a 1 µm scale bar at lower right.
Scanning electron micrograph of a cluster of Escherichia coli bacteria magnified 10,000 times; each rod-shaped cell is about one micrometre long (USDA ARS). · Photo by Eric Erbe, digital colorization by Christopher Pooley, both of USDA, ARS, EMU. · Public domain

5. Eukaryotic Cell Structure

Eukaryotic cells are distinguished by a membrane-bound nucleus and by numerous membrane-bound organelles, among them the endoplasmic reticulum, the Golgi apparatus, chloroplasts and mitochondria, and they carry several rod-shaped chromosomes. The nucleus is the largest and most prominent of the cell's organelles, measuring about 5 μm in diameter, and it houses the cell's DNA while directing the synthesis of ribosomes and proteins. It also acts as the command center of the cell, sending directions for the cell to grow, mature, divide or die.

Mitochondria are the powerhouses, or energy factories, of the cell, making ATP, the main energy-carrying molecule; they are found in almost all eukaryotic cells and generate most of the cell's supply of ATP from the breakdown of organic molecules. Chloroplasts are the sites of photosynthesis and are found only in plants and green algae. The endoplasmic reticulum is involved in the processing and transport of proteins and in the synthesis of lipids, and proteins are carried from it in small membrane vesicles to the Golgi apparatus for further processing and sorting.

Packed cells in glowing false colour: green filament bundles, yellow clouds of mitochondria and purple nuclei dotted with red.
Cultured HeLa cells with organelles revealed by fluorescent antibodies: microtubules in green, mitochondria in yellow, nucleoli in red and nuclear DNA in purple (GerryShaw). · GerryShaw · CC BY-SA 4.0

The cytoskeleton is a network of protein filaments extending throughout the cytoplasm, providing the cell with a structural framework and responsible for the movements of entire cells and for the positioning of organelles, including the movements of chromosomes during cell division. Lysosomes serve as the cell's garbage disposal, their enzymes breaking down proteins, polysaccharides, lipids, nucleic acids and worn-out organelles. Ribosomes are the structures responsible for protein synthesis, appearing as clusters or as single tiny dots floating freely in the cytoplasm.

Animal cells each have a centrosome and lysosomes, whereas most plant cells do not. Plant cells have a cell wall, chloroplasts and other specialized plastids, and a large central vacuole.

6. Cell Size and Scale

Cells and their structures span an enormous range of sizes, and the units used to describe them are correspondingly small. A typical human red blood cell is about eight micrometers in diameter, so that about 250 red blood cells could fit on a pinhead 12Biology 2e, 4.1 Studying Cells · OpenStax, Rice University ↗. Most bacterial cells are spherical, rod-shaped or spiral, with diameters of 1 to 10 μm; their DNA contents range from about 0.6 million to 5 million base pairs, enough to encode about 5000 different proteins 13The Origin and Evolution of Cells (The Cell: A Molecular Approach, 2nd edition) · NCBI Bookshelf / National Center for Biotechnology Information (Geoffrey M. Cooper) ↗.

Cell or structureSizeAdditional detail
Human red blood cellAbout 8 μm diameterAbout 250 could fit on a pinhead
Most bacterial cells1 to 10 μm diameterSpherical, rod-shaped or spiral
E. coliAbout 1 μm diameter and about 2 μm longRod-shaped
NucleusApproximately 5 μm diameter
Yeast Saccharomyces cerevisiaeAbout 6 μm diameter
Mycoplasmas, the smallest known cellsSpheres as small as 0.2 μm diameterTotal mass about 10−14 gram
Amoeba proteusLength can exceed 1 mmVolume more than 100,000 times that of E. coli
Representative sizes of cells and cell structures

7. Genomes and Ribosome Counts

Every cell carries its genetic material in the form of DNA, and the quantity of that DNA varies widely across cell types. In most bacteria, the genome spans roughly 0.6 million to 5 million base pairs, a complement sufficient to encode about 5000 different proteins 13. DNA is one of the four components common to all cells, alongside the plasma membrane, the cytoplasm, and the ribosomes that synthesize proteins 14Biology 2e, 4.2 Prokaryotic Cells · OpenStax, Rice University ↗. The scale of the genome is matched by the scale of the machinery that reads it: a single E. coli cell contains approximately 30,000 ribosomes.

The contrast between bacterial and human genomes is instructive. Human body (somatic) cells have 46 chromosomes, whereas human gametes — sperm or eggs — carry 23 chromosomes each 15Biology 2e, 10.1 Cell Division · OpenStax, Rice University ↗. Between these extremes sits the yeast Saccharomyces cerevisiae, whose genome comprises 12 million base pairs of DNA.

Cell or genomeDNA contentNotes
Bacterial genomeabout 0.6 million to 5 million base pairsenough to encode about 5000 different proteins
Yeast (Saccharomyces cerevisiae)12 million base pairs of DNA
Human somatic cells46 chromosomes
Human gametes23 chromosomessperm or eggs
E. coli ribosomesapproximately 30,000 per cell
DNA content and ribosome counts across selected cells and genomes.

8. Origins and Evolution

The spontaneous formation of organic molecules was first demonstrated experimentally in the 1950s, when Stanley Miller showed that electric sparks discharged into a mixture of hydrogen, methane and ammonia in the presence of water led to the formation of organic molecules including several amino acids.

A very early event in evolution appears to have been the divergence of three lines of descent from a common ancestor, giving rise to present-day archaebacteria, eubacteria and eukaryotes. Based upon the chloroplasts' ability to reproduce independently, the Russian botanist Konstantin Mereschkowski suggested in 1905 that chloroplasts may have originated from ancestral photosynthetic bacteria living symbiotically inside a eukaryotic cell.

9. The Cell Cycle

A cell reproduces by performing an orderly sequence of events in which it duplicates its contents and then divides in two, a cycle of duplication and division that is the essential mechanism by which all living things reproduce. In unicellular species such as bacteria and yeasts, each cell division produces a complete new organism, while in multicellular species long and complex sequences of cell divisions are required to produce a functioning organism.

The period between one M phase and the next is called interphase, and in most rapidly proliferating cells it is divided into three phases: S phase, in which DNA is replicated, and two gap phases (G1 and G2) that provide additional time for the cell to grow.

A field of cells with red rounded nuclei and green filament networks; at centre one cell shows a bright green spindle pulling its chromosomes apart; a 20 µm scale bar sits at lower right.
Fluorescence microscopy of LLC-PK1 cells with nuclei in red and the cytoskeleton in green; one cell is caught mid-division, its chromosomes separating on a bright spindle (ZEISS Microscopy). · ZEISS Microscopy from Germany · CC BY 2.0

10. Control of the Cell Cycle

The cell-cycle control system monitors progression through the cell cycle and delays later events until earlier events have been completed. When the system malfunctions, excessive cell divisions can result in cancer.

Progression through G1 and G2 is delayed by braking mechanisms if the DNA in the chromosomes is damaged by radiation or chemicals. Delays at these DNA damage checkpoints provide time for the damaged DNA to be repaired.

There are four classes of cyclins, each defined by the stage of the cell cycle at which they bind Cdks and function. Three of these classes are required in all eucaryotic cells.

11. Cell Division, Meiosis and Sexual Reproduction

Sexual reproduction requires the union of two specialized cells called gametes, each of which contains one set of chromosomes; when gametes unite they form a zygote that contains two sets of chromosomes. Meiosis is the nuclear division that forms haploid cells from diploid cells, and it employs many of the same cellular mechanisms as mitosis.It is a two-step process that reduces the chromosome number by half to form sperm and egg cells, and it also allows genetic variation through a process of gene shuffling.

Rows of roughly rectangular plant cells with brown-purple chromosomes; several cells show condensed chromosomes or two separating sets.
Micrograph of an onion root tip squash stained with acetocarmine; cells are visible at all the stages of mitosis, from prophase to telophase. · staticd · CC BY-SA 3.0

12. Timeline: from Hooke's Microscope to Super-Resolution Microscopy

  • 1665
    Hooke coins the term cell
    Robert Hooke described cork and other plant tissues in his Micrographia and introduced the term cell because the cellulose walls of dead cork cells reminded him of the blocks of cells occupied by monks.
  • From 1673
    Leeuwenhoek discovers the animalcules
    The Dutch microscopist Antonie van Leeuwenhoek began observing and describing blood cells, spermatozoa and a lively world of microscopic animalcules, opening the study of unicellular life.
  • 1831
    Brown describes the nucleus
    The Scottish botanist Robert Brown was the first to describe observations of nuclei, which he observed in plant cells.
  • 1838
    Schleiden and Schwann propose the cell theory
    The botanist Matthias Jakob Schleiden described plant tissues as composed of cells, and Theodor Schwann extended the cellular theory to animals, the first statement that all plants and animals are made up of cells.
  • 1839
    Cells as the elementary particles of organisms
    Schleiden and Schwann stated that cells are the elementary particles of organisms in both plants and animals, recognizing that some organisms are unicellular and others multicellular, in Schwann's Mikroskopische Untersuchungen.
  • 1852
    Remak shows cells divide
    Robert Remak, a neurologist and embryologist, published convincing evidence that cells are derived from other cells as a result of cell division.
  • 1855
    Virchow's Cellular Pathology
    Rudolf Virchow published an editorial essay entitled Cellular Pathology, popularizing the Latin phrase omnis cellula a cellula, all cells arise from cells.
  • 1905
    The first endosymbiotic hypothesis
    The Russian botanist Konstantin Mereschkowski suggested that chloroplasts may have originated from ancestral photosynthetic bacteria living symbiotically inside a eukaryotic cell.
  • 1933
    The first electron microscope
    Ernst Ruska discovered that a magnetic coil could be used as a lens for electron beams and developed the first electron microscope, capturing images of objects far smaller than the wavelength of light.
  • c. 1953
    Miller makes organic molecules from inorganic gases
    Stanley Miller showed that electric sparks discharged into a mixture of hydrogen, methane and ammonia in the presence of water led to the formation of organic molecules, including several amino acids.
  • 1967
    Margulis revives the endosymbiotic hypothesis
    Lynn Margulis, an American geneticist, published her ideas on the endosymbiotic origin of mitochondria and chloroplasts, reviving a hypothesis that became central to modern cell biology.
  • 2014
    Super-resolved fluorescence microscopy
    The Nobel Prize in Chemistry 2014 was awarded to Eric Betzig, Stefan W. Hell and William E. Moerner for the development of super-resolved fluorescence microscopy, allowing cells to be studied at the nanoscale.

13. Frequently asked questions

How long does the cell cycle take in a typical mammalian cell?

In a typical human cell proliferating in culture, interphase might occupy 23 hours of a 24 hour cycle, with the G1, S and G2 phases together making up that interphase.20An Overview of the Cell Cycle (Molecular Biology of the Cell, 4th edition) · NCBI Bookshelf / National Center for Biotechnology Information (Alberts et al.) ↗ DNA duplication occurs during S phase, which requires 10 to 12 hours and occupies about half of the cell-cycle time in a typical mammalian cell.20 M phase, by contrast, lasts less than an hour.20

What happens to a cell that is not ready to divide?

If extracellular conditions are unfavorable, cells delay progress through G1 and may even enter a specialized resting state known as G0, in which they can remain for days, weeks or even years before resuming proliferation.20

What are the checkpoints that guard the cell cycle?

Entry into mitosis is prevented when DNA replication is not complete, and chromosome separation in mitosis is delayed if some chromosomes are not properly attached to the mitotic spindle.17Components of the Cell-Cycle Control System (Molecular Biology of the Cell, 4th edition) · NCBI Bookshelf / National Center for Biotechnology Information (Alberts et al.) ↗

What actually drives the cell cycle forward?

At the heart of the cell-cycle control system is a family of protein kinases known as cyclin-dependent kinases, whose activity rises and falls as the cell progresses through the cycle.17 The system operates much like the control system of an automatic clothes-washing machine, with a central controller triggering each process in a set sequence.17

How does a cell avoid copying its DNA twice in one cycle?

The activation of S-Cdk in late G1 pulls the trigger and initiates DNA replication, and S-Cdk also helps to prevent rereplication by causing the Cdc6 protein to dissociate from ORC after an origin has fired.21Intracellular Control of Cell-Cycle Events (Molecular Biology of the Cell, 4th edition) · NCBI Bookshelf / National Center for Biotechnology Information (Alberts et al.) ↗ Early clues about the regulation of S phase came from studies in which human cells at various cell-cycle stages were fused to form single cells with two nuclei.21

What happens to the cell's internal architecture during mitosis?

During mitosis the chromosomes condense, the nuclear envelope breaks down, the endoplasmic reticulum and Golgi apparatus reorganize, and the cytoskeleton radically reorganizes to segregate the replicated chromosomes and divide the cell in two.16The Mechanics of Cell Division (Molecular Biology of the Cell, 4th edition, Chapter 18) · NCBI Bookshelf / National Center for Biotechnology Information (Alberts et al.) ↗ Targeted protein degradation by the anaphase-promoting complex initiates the separation and segregation of the replicated chromosomes and inactivates M-Cdk at the end of mitosis.16

References (22)

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