Cell

Cell (biology

Allium cells in different phases of the cell cycle

Cells in culture, stained for keratin (red) and DNA (green)

Drawing of the structure of cork as it appeared under the microscope to Robert Hooke from Micrographia, which is the origin of the word "cell" being used to describe the smallest unit of a living organism

The cell is the functional basic unit of life. It was discovered by Robert Hooke and is the functional unit of all known living organisms. It is the smallest unit of life that is classified as a living thing, and is often called the building block of life.^[1] Organisms can be classified as unicellular (consisting of a single cell; including most bacteria) or multicellular (including plants and animals). Humans contain about 100 trillion cells; a typical cell size is 10 µm and a typical cell mass is 1 nanogram. The human cell extrema are: largest - anterior horn in the spinal cord (135 μ vs. 120 μ for the ovum), longest - pseudounipolar cells which reach from extremities, including the toes to the lower brain stem, and smallest - granule cells in the cerebellum, at 4 µ.^[2] The largest known cells are unfertilised ostrich egg cells, which weigh 3.3 pounds.^[3][4]
In 1835, before the final cell theory was developed, Jan Evangelista Purkyně observed small "granules" while looking at the plant tissue through a microscope. The 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 all cells come from preexisting cells, that vital functions of an organism occur within cells, and that all cells contain the hereditary information necessary for regulating cell functions and for transmitting information to the next generation of cells.^[5]
The word cell comes from the Latin cellula, meaning "a small room". The descriptive term for the smallest living biological structure was coined by Robert Hooke in a book he published in 1665 when he compared the cork cells he saw through his microscope to the small rooms monks lived in.^[6]

 

Anatomy

The cells of eukaryotes (left) and prokaryotes (right)

There are two types of cells: eukaryotic and prokaryotic. Prokaryotic cells are usually independent, while eukaryotic cells are often found in multicellular organisms.

Table 1: Comparison of features of prokaryotic and eukaryotic cells

	Prokaryotes	Eukaryotes
Typical organisms	bacteria, archaea	protists, fungi, plants, animals
Typical size	~ 1–10 µm	~ 10–100 µm (sperm cells, apart from the tail, are smaller)
Type of nucleus	nucleoid region; no real nucleus	real nucleus with double membrane
DNA	circular (usually)	linear molecules (chromosomes) with histone proteins
RNA-/protein-synthesis	coupled in cytoplasm	RNA-synthesis inside the nucleus protein synthesis in cytoplasm
Ribosomes	50S+30S	60S+40S
Cytoplasmatic 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 (though some lack mitochondria)
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

Prokaryotic cells

Main article: Prokaryote

Diagram of a typical prokaryotic cell

The prokaryote cell is simpler, and therefore smaller, than a eukaryote cell, lacking a nucleus and most of the other organelles of eukaryotes. There are two kinds of prokaryotes: bacteria and archaea; these share a similar structure.
Nuclear material of prokaryotic cell consist of a single chromosome that is in direct contact with cytoplasm. Here, the undefined nuclear region in the cytoplasm is called nucleoid.
A prokaryotic cell has three architectural regions:

• On the outside, flagella and pili project from the cell's surface. These are structures (not present in all prokaryotes) made of proteins that facilitate movement and communication between cells;
• Enclosing the cell is the cell envelope – generally consisting of a cell wall covering a plasma membrane though some bacteria also have a further covering layer called a capsule. The envelope gives rigidity to the cell and separates the interior of the cell from its environment, serving as a protective filter. Though most prokaryotes have a cell wall, there are exceptions such as Mycoplasma (bacteria) and Thermoplasma (archaea). The cell wall consists of peptidoglycan in bacteria, and acts as an additional barrier against exterior forces. It also prevents the cell from expanding and finally bursting (cytolysis) from osmotic pressure against a hypotonic environment. Some eukaryote cells (plant cells and fungi cells) also have a cell wall;
• Inside the cell is the cytoplasmic region that contains the cell genome (DNA) and ribosomes and various sorts of inclusions. A prokaryotic chromosome is usually a circular molecule (an exception is that of the bacterium Borrelia burgdorferi, which causes Lyme disease). Though not forming a nucleus, the DNA is condensed in a nucleoid. Prokaryotes can carry extrachromosomal DNA elements called plasmids, which are usually circular. Plasmids enable additional functions, such as antibiotic resistance.

Eukaryotic cells

Main article: Eukaryote

Plants, animals, fungi, slime moulds, protozoa, & algae are all Eukaryotic. These cells are about 15 times wider than a typical prokaryote and can be as much as 1000 times greater in volume. The major difference between prokaryotes and eukaryotes is that eukaryotic cells contain membrane-bound compartments in which specific metabolic activities take place. Most important among these is a cell nucleus, a membrane-delineated compartment that houses the eukaryotic cell's DNA. This nucleus gives the eukaryote its name, which means "true nucleus." Other differences include:

• The plasma membrane resembles that of prokaryotes in function, with minor differences in the setup. Cell walls may or may not be present.
• The eukaryotic DNA is organized in one or more linear molecules, called chromosomes, which are associated with histone proteins. All chromosomal DNA is stored in the cell nucleus, separated from the cytoplasm by a membrane. Some eukaryotic organelles such as mitochondria also contain some DNA.
• Many eukaryotic cells are ciliated with primary cilia. Primary cilia play important roles in chemosensation, mechanosensation, and thermosensation. Cilia may thus be "viewed as sensory cellular antennae that coordinate a large number of cellular signaling pathways, sometimes coupling the signaling to ciliary motility or alternatively to cell division and differentiation."^[7]
• Eukaryotes can move using motile cilia or flagella. The flagella are more complex than those of prokaryotes.

Structure of a typical animal cell

Structure of a typical plant cell

Table 2: Comparison of structures between animal and plant cells

	Typical animal cell	Typical plant cell
Organelles	Nucleus Nucleolus (within nucleus) Rough endoplasmic reticulum (ER) Smooth ER Ribosomes Cytoskeleton Golgi apparatus Cytoplasm Mitochondria Vesicles Lysosomes Centrosome Centrioles	Nucleus Nucleolus (within nucleus) Rough ER Smooth ER Ribosomes Cytoskeleton Golgi apparatus (dictiosomes) Cytoplasm Mitochondria Plastids and its derivatives Vacuole(s) Cell wall

Subcellular components

All cells, whether prokaryotic or eukaryotic, have a membrane that envelops the cell, separates its interior from its environment, regulates what moves in and out (selectively permeable), and maintains the electric potential of the cell. Inside the membrane, a salty cytoplasm takes up most of the cell volume. 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 components of the cell, then briefly describe their function.

Membrane

Main article: Cell membrane

The cytoplasm of a cell is surrounded by a cell membrane or plasma membrane. The plasma membrane in plants and prokaryotes 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 lipids (hydrophobic fat-like molecules) and hydrophilic phosphorus molecules. Hence, the layer is called a phospholipid bilayer. It may also be called a fluid mosaic membrane. Embedded within this membrane is a variety of protein molecules that act as channels and pumps that move different molecules into and out of the cell. The membrane is said to be 'semi-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

Bovine Pulmonary Artery Endothelial cell: nuclei stained blue, mitochondria stained red, and F-actin, an important component in microfilaments, stained green. Cell imaged on a fluorescent microscope.

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 microfilaments, intermediate filaments and microtubules. There is a great number of proteins associated with them, each controlling a cell's structure by directing, bundling, and aligning filaments. The prokaryotic cytoskeleton is less well-studied but is involved in the maintenance of cell shape, polarity and cytokinesis.^[8]

Genetic material

Two different kinds of genetic material exist: deoxyribonucleic acid (DNA) and ribonucleic acid (RNA). Most organisms use DNA for their long-term information storage, but some viruses (e.g., retroviruses) have RNA as their genetic material. The biological information contained in an organism is encoded in its DNA or RNA sequence. RNA is also used for information transport (e.g., mRNA) and enzymatic functions (e.g., ribosomal RNA) in organisms that use DNA for the genetic code itself. Transfer RNA (tRNA) molecules are used to add amino acids during protein translation.
Prokaryotic genetic material is organized in a simple circular DNA molecule (the bacterial chromosome) in the nucleoid region of the cytoplasm. Eukaryotic genetic material is divided into different, 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 23 pairs of linear DNA molecules called chromosomes. The mitochondrial genome is a circular DNA molecule distinct from the nuclear DNA. Although the mitochondrial DNA is very small compared to nuclear chromosomes, 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

The human body contains many different organs, such as the heart, lung, and kidney, with each organ performing a different function. Cells also have a set of "little organs," called organelles, that are adapted and/or specialized for carrying out one or more vital functions. Both eukaryotic and prokaryotic cells have organelles but organelles in eukaryotes are generally more complex and may be 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, peroxisomes and lysosomes) can be numerous (hundreds to thousands). The cytosol is the gelatinous fluid that fills the cell and surrounds the organelles.

Cell nucleus – eukaryotes only - a cell's information center  The cell nucleus is the most conspicuous organelle found in a eukaryotic cell. It houses the cell's chromosomes, and is the place where almost all DNA replication and RNA synthesis (transcription) occur. The nucleus is spherical and separated from the cytoplasm by a double membrane called the nuclear envelope. The nuclear envelope isolates and protects a cell's DNA from various molecules that could accidentally damage its structure or interfere with its processing. During processing, DNA is transcribed, or copied into a special RNA, called messenger RNA (mRNA). This mRNA is then transported out of the nucleus, where it is translated into a specific protein molecule. The nucleolus is a specialized region within the nucleus where ribosome subunits are assembled. In prokaryotes, DNA processing takes place in the cytoplasm.	Diagram of a cell nucleus
Mitochondria and Chloroplasts – eukaryotes only - the power generators  Mitochondria are self-replicating organelles that occur in various numbers, shapes, and sizes in the cytoplasm of all eukaryotic cells. Mitochondria play a critical role in generating energy in the eukaryotic cell. Mitochondria generate the cell's energy by oxidative phosphorylation, using oxygen to release energy stored in cellular nutrients (typically pertaining to glucose) to generate ATP. Mitochondria multiply by splitting in two. Respiration occurs in the cell mitochondria. Organelles that are modified chloroplasts are broadly called plastids, and are involved in energy storage through photosynthesis, which uses solar energy to generate carbohydrates and oxygen from carbon dioxide and water.[citation needed] Mitochondria and chloroplasts each contain their own genome, which is separate and distinct from the nuclear genome of a cell. Both organelles contain this DNA in circular plasmids, much like prokaryotic cells, strongly supporting the evolutionary theory of endosymbiosis; since these organelles contain their own genomes and have other similarities to prokaryotes, they are thought to have developed through a symbiotic relationship after being engulfed by a primitive cell.[citation needed]
Endoplasmic reticulum – eukaryotes only  The endoplasmic reticulum (ER) is the transport network for molecules targeted for certain modifications and specific destinations, as compared to molecules that float freely in the cytoplasm. The ER has two forms: the rough ER, which has ribosomes on its surface and secretes proteins into the cytoplasm, and the smooth ER, which lacks them. Smooth ER plays a role in calcium sequestration and release.
Golgi apparatus – eukaryotes only  The primary function of the Golgi apparatus is to process and package the macromolecules such as proteins and lipids that are synthesized by the cell. It is particularly important in the processing of proteins for secretion. The Golgi apparatus forms a part of the endomembrane system of eukaryotic cells. Vesicles that enter the Golgi apparatus are processed in a cis to trans direction, meaning they coalesce on the cis side of the apparatus and after processing pinch off on the opposite (trans) side to form a new vesicle in the animal cell.[citation needed]	Diagram of an endomembrane system
Ribosomes  The ribosome is a large complex of RNA and protein molecules. They each consist of two subunits, and act as an assembly line where RNA from the nucleus is used to synthesise proteins from amino acids. Ribosomes can be found either floating freely or bound to a membrane (the rough endoplasmatic reticulum in eukaryotes, or the cell membrane in prokaryotes).[9]
Lysosomes and Peroxisomes – eukaryotes only  Lysosomes contain digestive enzymes (acid hydrolases). They digest excess or worn-out organelles, food particles, and engulfed viruses or bacteria. Peroxisomes have enzymes that rid the cell of toxic peroxides. The cell could not house these destructive enzymes if they were not contained in a membrane-bound system. These organelles are often called a "suicide bag" because of their ability to detonate and destroy the cell.[citation needed]
Centrosome – the cytoskeleton organiser  The centrosome produces the microtubules of a cell – a key component of the cytoskeleton. It directs the transport through the ER and the Golgi apparatus. Centrosomes are composed of two centrioles, which separate during cell division and help in the formation of the mitotic spindle. A single centrosome is present in the animal cells. They are also found in some fungi and algae cells.[citation needed]
Vacuoles  Vacuoles store food and waste. Some vacuoles store extra water. They are often described as liquid filled space and are surrounded by a membrane. Some cells, most notably Amoeba, have contractile vacuoles, which can pump water out of the cell if there is too much water. The vacuoles of eukaryotic cells are usually larger in those of plants than animals.

Structures outside the cell wall

Capsule

A gelatinous capsule is present in some bacteria outside the cell wall. The capsule may be polysaccharide as in pneumococci, meningococci or polypeptide as Bacillus anthracis or hyaluronic acid as in streptococci.^{[citation needed]} Capsules are not marked by ordinary stain and can be detected by special stain. The capsule is antigenic. The capsule has antiphagocytic function so it determines the virulence of many bacteria. It also plays a role in attachment of the organism to mucous membranes.^{[citation needed]}

Flagella

Flagella are the organelles of cellular mobility. They arise from cytoplasm and extrude through the cell wall. They are long and thick thread-like appendages, protein in nature. Are most commonly found in bacteria cells but are found in animal cells as well.

Fimbriae (pili)

They are short and thin hair like filaments, formed of protein called pilin (antigenic). Fimbriae are responsible for attachment of bacteria to specific receptors of human cell (adherence). There are special types of pili called (sex pili) involved in conjunction.^{[citation needed]}

Functions

Growth and metabolism

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 a less chemically complex sugar molecule called glucose. Once inside the cell, glucose is broken down to make adenosine triphosphate (ATP), a form of energy, through two different pathways.
The first pathway, glycolysis, requires no oxygen and is referred to as anaerobic metabolism. Each reaction is designed to produce some hydrogen ions that can then be used to make energy packets (ATP). In prokaryotes, glycolysis is the only method used for converting energy.
The second pathway, called the Krebs cycle, or citric acid cycle, occurs inside the mitochondria and can generate enough ATP to run all the cell functions.

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.

Creation

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. 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, is required every time a cell divides. Replication, like all cellular activities, requires specialized proteins for carrying out the job.

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.

Movement or motility

Cells can move during many processes: such as wound healing, the immune response and cancer metastasis. For wound healing to occur, white blood cells and cells that ingest bacteria move to the wound site to kill the microorganisms that cause infection.
At the same time fibroblasts (connective tissue cells) move there to remodel damaged structures. In the case of tumor development, cells from a primary tumor move away and spread to other parts of the body. Cell motility involves many receptors, crosslinking, bundling, binding, adhesion, motor and other proteins.^[10] 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.^[11][12]

Evolution

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

Further information: Abiogenesis

There are several theories about the origin of small molecules that could lead to life in an early Earth. One is that they came from meteorites (see Murchison meteorite). Another is that they were created at deep-sea vents. A third is that they were synthesized by lightning in a reducing atmosphere (see Miller–Urey experiment); although it is not clear if Earth had such an atmosphere. There are essentially no experimental data defining what the first self-replicating forms were. RNA is generally assumed 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, like clay or peptide nucleic acid.^[13]
Cells emerged at least 4.0–4.3 billion years ago. The current belief is that these cells were heterotrophs. An important characteristic of cells is the cell membrane, composed of a bilayer of lipids. 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.^[14]

Origin of eukaryotic cells

The eukaryotic cell seems to have evolved from a symbiotic community of prokaryotic cells. DNA-bearing organelles like the mitochondria and the chloroplasts are almost certainly what remains of ancient symbiotic oxygen-breathing proteobacteria and cyanobacteria, respectively, where the rest of the cell appears derived from an ancestral archaean prokaryote cell—an idea called the endosymbiotic theory.
There is still considerable debate about whether organelles like the hydrogenosome predated the origin of mitochondria, or viceversa: see the hydrogen hypothesis for the origin of eukaryotic cells.
Sex, as the stereotyped choreography of meiosis and syngamy that persists in nearly all extant eukaryotes, may have played a role in the transition from prokaryotes to eukaryotes. An 'origin of sex as vaccination' theory suggests that the eukaryote genome accreted from prokaryan parasite genomes in numerous rounds of lateral gene transfer. Sex-as-syngamy (fusion sex) arose when infected hosts began swapping nuclearized genomes containing co-evolved, vertically transmitted symbionts that conveyed protection against horizontal infection by more virulent symbionts.^[15]

History

• 1632–1723: Antonie van Leeuwenhoek teaches himself to grind lenses, builds a microscope and draws protozoa, such as Vorticella from rain water, and bacteria from his own mouth.
• 1665: Robert Hooke discovers cells in cork, then in living plant tissue using an early microscope.^[6]
• 1839: Theodor Schwann and Matthias Jakob Schleiden elucidate the principle that plants and animals are made of cells, concluding that cells are a common unit of structure and development, and thus founding the cell theory.
• The belief that life forms can occur spontaneously (generatio spontanea) is contradicted by Louis Pasteur (1822–1895) (although Francesco Redi had performed an experiment in 1668 that suggested the same conclusion).
• 1855: Rudolf Virchow states that cells always emerge from cell divisions (omnis cellula ex cellula).
• 1931: Ernst Ruska builds first transmission electron microscope (TEM) at the University of Berlin. By 1935, he has built an EM with twice the resolution of a light microscope, revealing previously unresolvable organelles.
• 1953: Watson and Crick made their first announcement on the double-helix structure for DNA on February 28.
• 1981: Lynn Margulis published Symbiosis in Cell Evolution detailing the endosymbiotic theory.

Light

Light or visible light is electromagnetic radiation that is visible to the human eye, and is responsible for the sense of sight.^[1] Visible light haswavelength in a range from about 380 nanometres to about 740 nm, with a frequency range of about 405 THz to 790 THz. In physics, the term lightsometimes refers to electromagnetic radiation of any wavelength, whether visible or not.^[2][3]

Primary properties of light are intensity, propagation direction, frequency or wavelength spectrum, and polarisation, while its speed in a vacuum, 299,792,458 meters per second (about 300,000 kilometers per second), is one of the fundamental constants of nature.

Light, which is emitted and absorbed in tiny "packets" called photons, exhibits properties of both waves and particles. This property is referred to as thewave–particle duality. The study of light, known as optics, is an important research area in modern physics.

Speed of light

Main article: Speed of light

The speed of light in a vacuum is defined to be exactly 299,792,458 m/s (approximately 186,282 miles per second). The fixed value of the speed of light in SI units results from the fact that the metre is now defined in terms of the speed of light.

Different physicists have attempted to measure the speed of light throughout history. Galileo attempted to measure the speed of light in the seventeenth century. An early experiment to measure the speed of light was conducted by Ole Rømer, a Danish physicist, in 1676. Using a telescope, Rømer observed the motions of Jupiter and one of its moons, Io. Noting discrepancies in the apparent period of Io's orbit, he calculated that light takes about 22 minutes to traverse the diameter of Earth's orbit.^[4] Unfortunately, its size was not known at that time. If Rømer had known the diameter of the Earth's orbit, he would have calculated a speed of 227,000,000 m/s.

Another, more accurate, measurement of the speed of light was performed in Europe by Hippolyte Fizeau in 1849. Fizeau directed a beam of light at a mirror several kilometers away. A rotating cog wheel was placed in the path of the light beam as it traveled from the source, to the mirror and then returned to its origin. Fizeau found that at a certain rate of rotation, the beam would pass through one gap in the wheel on the way out and the next gap on the way back. Knowing the distance to the mirror, the number of teeth on the wheel, and the rate of rotation, Fizeau was able to calculate the speed of light as 313,000,000 m/s.

Léon Foucault used an experiment which used rotating mirrors to obtain a value of 298,000,000 m/s in 1862. Albert A. Michelson conducted experiments on the speed of light from 1877 until his death in 1931. He refined Foucault's methods in 1926 using improved rotating mirrors to measure the time it took light to make a round trip from Mt. Wilson to Mt. San Antonio in California. The precise measurements yielded a speed of 299,796,000 m/s.

Two independent teams of physicists were able to bring light to a complete standstill by passing it through a Bose-Einstein Condensate of the element rubidium, one team led by Dr. Lene Vestergaard Hau of Harvard University and the Rowland Institute for Science in Cambridge, Mass., and the other by Dr. Ronald L. Walsworth and Dr. Mikhail D. Lukin of the Harvard-Smithsonian Center for Astrophysics, also in Cambridge.^[5]

Electromagnetic spectrum

Main article: Electromagnetic spectrum

Electromagnetic spectrum with light highlighted

Generally, EM radiation (the designation 'radiation' excludes static electric and magnetic and near fields) is classified by wavelength into radio, microwave, infrared, the visible region we perceive as light, ultraviolet, X-rays and gamma rays.

The behaviour of EM radiation depends on its wavelength. Higher frequencies have shorter wavelengths, and lower frequencies have longer wavelengths. When EM radiation interacts with single atoms and molecules, its behaviour depends on the amount of energy per quantum it carries.

Optics

Main article: Optics

The study of light and the interaction of light and matter is termed optics. The observation and study of optical phenomena such as rainbows and the aurora borealis offer many clues as to the nature of light as well as much enjoyment.

Refraction

Main article: Refraction

An example of refraction of light. The straw appears bent, because of refraction of light as it enters liquid from air.

Refraction is the bending of light rays when passing through a surface between one transparent material and another. It is described by Snell's Law:

where θ₁ is the angle between the ray and the surface normal in the first medium, θ₂ is the angle between the ray and the surface normal in the second medium, and n₁ and n₂ are the indices of refraction, n = 1 in a vacuum and n > 1 in a transparent substance.

When a beam of light crosses the boundary between a vacuum and another medium, or between two different media, the wavelength of the light changes, but the frequency remains constant. If the beam of light is not orthogonal (or rather normal) to the boundary, the change in wavelength results in a change in the direction of the beam. This change of direction is known as refraction.

The refractive quality of lenses is frequently used to manipulate light in order to change the apparent size of images. Magnifying glasses, spectacles,contact lenses, microscopes and refracting telescopes are all examples of this manipulation.

Light sources

See also: List of light sources

A cloud illuminated by sunlight

There are many sources of light. The most common light sources are thermal: a body at a given temperature emits a characteristic spectrum of black-body radiation. Examples include sunlight (the radiation emitted by the chromosphere of the Sun at around 6,000 Kelvin peaks in the visible region of the electromagnetic spectrum when plotted in wavelength units ^[6] and roughly 40% of sunlight is visible), incandescent light bulbs (which emit only around 10% of their energy as visible light and the remainder as infrared), and glowing solid particles in flames. The peak of the blackbody spectrum is in the infrared for relatively cool objects like human beings. As the temperature increases, the peak shifts to shorter wavelengths, producing first a red glow, then a white one, and finally a blue colour as the peak moves out of the visible part of the spectrum and into the ultraviolet. These colours can be seen when metal is heated to "red hot" or "white hot". Blue thermal emission is not often seen. The commonly seen blue colour in a gas flame or a welder'storch is in fact due to molecular emission, notably by CH radicals (emitting a wavelength band around 425 nm).

Atoms emit and absorb light at characteristic energies. This produces "emission lines" in the spectrum of each atom. Emission can be spontaneous, as in light-emitting diodes, gas discharge lamps (such as neon lamps and neon signs, mercury-vapor lamps, etc.), and flames (light from the hot gas itself—so, for example, sodium in a gas flame emits characteristic yellow light). Emission can also be stimulated, as in a laser or a microwave maser.

Deceleration of a free charged particle, such as an electron, can produce visible radiation: cyclotron radiation, synchrotron radiation, and bremsstrahlungradiation are all examples of this. Particles moving through a medium faster than the speed of light in that medium can produce visible Cherenkov radiation.

Certain chemicals produce visible radiation by chemoluminescence. In living things, this process is called bioluminescence. For example, firefliesproduce light by this means, and boats moving through water can disturb plankton which produce a glowing wake.

Certain substances produce light when they are illuminated by more energetic radiation, a process known as fluorescence. Some substances emit light slowly after excitation by more energetic radiation. This is known as phosphorescence.

Phosphorescent materials can also be excited by bombarding them with subatomic particles. Cathodoluminescence is one example. This mechanism is used in cathode ray tube television sets andcomputer monitors.

A city illuminated by artificial lighting

Certain other mechanisms can produce light:

• scintillation
• electroluminescence
• sonoluminescence
• triboluminescence
• Cherenkov radiation

When the concept of light is intended to include very-high-energy photons (gamma rays), additional generation mechanisms include:

• Radioactive decay
• Particle–antiparticle annihilation

Units and measures

Main articles: Photometry (optics) and Radiometry

Light is measured with two main alternative sets of units: radiometry consists of measurements of light power at all wavelengths, while photometry measures light with wavelength weighted with respect to a standardised model of human brightness perception. Photometry is useful, for example, to quantify Illumination (lighting) intended for human use. The SI units for both systems are summarised in the following tables.

Table 1. SI radiometry unitsv · d · e

Quantity	Symbol[nb 1]	SI unit	Symbol	Dimension	Notes
Radiant energy	Qe[nb 2]	joule	J	M⋅L2⋅T−2	energy
Radiant flux	Φe[nb 2]	watt	W	M⋅L2⋅T−3	radiant energy per unit time, also called radiant power.
Spectral power	Φeλ[nb 2][nb 3]	watt per metre	W⋅m−1	M⋅L⋅T−3	radiant power per wavelength.
Radiant intensity	Ie	watt per steradian	W⋅sr−1	M⋅L2⋅T−3	power per unit solid angle.
Spectral intensity	Ieλ[nb 3]	watt per steradian per metre	W⋅sr−1⋅m−1	M⋅L⋅T−3	radiant intensity per wavelength.
Radiance	Le	watt per steradian persquare metre	W⋅sr−1⋅m−2	M⋅T−3	power per unit solid angle per unit projected source area. confusingly called "intensity" in some other fields of study.
Spectral radiance	Leλ[nb 3] or Leν[nb 4]	watt per steradian per metre3 or watt per steradian per square metre per hertz	W⋅sr−1⋅m−3 or W⋅sr−1⋅m−2⋅Hz−1	M⋅L−1⋅T−3 or M⋅T−2	commonly measured in W⋅sr−1⋅m−2⋅nm−1 with surface area and either wavelength orfrequency.
Irradiance	Ee[nb 2]	watt per square metre	W⋅m−2	M⋅T−3	power incident on a surface, also called radiant flux density. sometimes confusingly called "intensity" as well.
Spectral irradiance	Eeλ[nb 3] or Eeν[nb 4]	watt per metre3 or watt per square metre per hertz	W⋅m−3 or W⋅m−2⋅Hz−1	M⋅L−1⋅T−3 or M⋅T−2	commonly measured in W⋅m−2⋅nm−1 or 10−22W⋅m−2⋅Hz−1, known as solar flux unit.[nb 5]
Radiant exitance / Radiant emittance	Me[nb 2]	watt per square metre	W⋅m−2	M⋅T−3	power emitted from a surface.
Radiosity	Je orJeλ[nb 3]	watt per square metre	W⋅m−2	M⋅T−3	emitted plus reflected power leaving a surface.
Radiant exposure	He	joule per square metre	J⋅m−2	M⋅T−2
Radiant energy density	ωe	joule per metre3	J⋅m−3	M⋅L−1⋅T−2
See also: SI · Radiometry · Photometry · (Compare)

Table 2. SI photometry unitsv · d · e

Quantity	Symbol[nb 6]	SI unit	Symbol	Dimension	Notes
Luminous energy	Qv [nb 7]	lumen second	lm⋅s	T⋅J	units are sometimes called talbots
Luminous flux	Φv [nb 7]	lumen (= cd⋅sr)	lm	J	also called luminous power
Luminous intensity	Iv	candela (= lm/sr)	cd	J [nb 8]	an SI base unit, luminous flux per unit solid angle
Luminance	Lv	candela per square metre	cd/m2	L−2⋅J	units are sometimes called nits
Illuminance	Ev	lux (= lm/m2)	lx	L−2⋅J	used for light incident on a surface
Luminous emittance	Mv	lux (= lm/m2)	lx	L−2⋅J	used for light emitted from a surface
Luminous exposure	Hv	lux second	lx⋅s	L−2⋅T⋅J
Luminous energy density	ωv	lumen second per metre3	lm⋅s⋅m−3	L−3⋅T⋅J
Luminous efficacy	η [nb 7]	lumen per watt	lm/W	M−1⋅L−2⋅T3⋅J	ratio of luminous flux to radiant flux
Luminous efficiency	V			1	also called luminous coefficient
See also: SI · Photometry · Radiometry · (Compare)

The photometry units are different from most systems of physical units in that they take into account how the human eye responds to light. The cone cells in the human eye are of three types which respond differently across the visible spectrum, and the cumulative response peaks at a wavelength of around 555 nm. Therefore, two sources of light which produce the same intensity (W/m²) of visible light do not necessarily appear equally bright. The photometry units are designed to take this into account, and therefore are a better representation of how "bright" a light appears to be than raw intensity. They relate to raw power by a quantity called luminous efficacy, and are used for purposes like determining how to best achieve sufficient illumination for various tasks in indoor and outdoor settings. The illumination measured by a photocell sensor does not necessarily correspond to what is perceived by the human eye, and without filters which may be costly, photocells and charge-coupled devices (CCD) tend to respond to some infrared, ultraviolet or both.

Light pressure

Main article: Radiation pressure

Light exerts physical pressure on objects in its path, a phenomenon which can be deduced by Maxwell's equations, but can be more easily explained by the particle nature of light: photons strike and transfer their momentum. Light pressure is equal to the power of the light beam divided by c, the speed of light.  Due to the magnitude of c, the effect of light pressure is negligible for everyday objects. For example, a one-milliwatt laser pointer exerts a force of about 3.3 piconewtons on the object being illuminated; thus, one could lift a U. S. penny with laser pointers, but doing so would require about 30 billion 1-mW laser pointers.^[7]  However, in nanometer-scale applications such as NEMS, the effect of light pressure is more pronounced, and exploiting light pressure to drive NEMS mechanisms and to flip nanometer-scale physical switches in integrated circuits is an active area of research.^[8]

At larger scales, light pressure can cause asteroids to spin faster,^[9] acting on their irregular shapes as on the vanes of a windmill.  The possibility to make solar sails that would accelerate spaceships in space is also under investigation.^[10][11]

Although the motion of the Crookes radiometer was originally attributed to light pressure, this interpretation is incorrect; the characteristic Crookes rotation is the result of a partial vacuum.^[12] This should not be confused with the Nichols radiometer, in which the motion is directly caused by light pressure.^[13]

Historical theories about light, in chronological order

Hindu and Buddhist theories

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In ancient India, the Hindu schools of Samkhya and Vaisheshika, from around the 6th–5th century BC, developed theories on light. According to the Samkhya school, light is one of the five fundamental "subtle" elements (tanmatra) out of which emerge the gross elements. The atomicity of these elements is not specifically mentioned and it appears that they were actually taken to be continuous.

On the other hand, the Vaisheshika school gives an atomic theory of the physical world on the non-atomic ground of ether, space and time. (See Indian atomism.) The basic atoms are those of earth (prthivı), water (pani), fire (agni), and air (vayu), that should not be confused with the ordinary meaning of these terms. These atoms are taken to form binary molecules that combine further to form larger molecules. Motion is defined in terms of the movement of the physical atoms and it appears that it is taken to be non-instantaneous. Light rays are taken to be a stream of high velocity of tejas (fire) atoms. The particles of light can exhibit different characteristics depending on the speed and the arrangements of the tejas atoms. Around the first century BC, the Vishnu Purana refers to sunlight as "the seven rays of the sun".

The Indian Buddhists, such as Dignāga in the 5th century and Dharmakirti in the 7th century, developed a type of atomism that is a philosophy about reality being composed of atomic entities that are momentary flashes of light or energy. They viewed light as being an atomic entity equivalent to energy, similar to the modern concept of photons, though they also viewed all matter as being composed of these light/energy particles.

It is written in the Rigveda that light consists of three primary colours. "Mixing the three colours, ye have produced all the objects of sight!"^[14]

Greek and Hellenistic theories

In the fifth century BC, Empedocles postulated that everything was composed of four elements; fire, air, earth and water. He believed that Aphrodite made the human eye out of the four elements and that she lit the fire in the eye which shone out from the eye making sight possible. If this were true, then one could see during the night just as well as during the day, so Empedocles postulated an interaction between rays from the eyes and rays from a source such as the sun.

In about 300 BC, Euclid wrote Optica, in which he studied the properties of light. Euclid postulated that light travelled in straight lines and he described the laws of reflection and studied them mathematically. He questioned that sight is the result of a beam from the eye, for he asks how one sees the stars immediately, if one closes one's eyes, then opens them at night. Of course if the beam from the eye travels infinitely fast this is not a problem.

In 55 BC, Lucretius, a Roman who carried on the ideas of earlier Greek atomists, wrote:

"The light & heat of the sun; these are composed of minute atoms which, when they are shoved off, lose no time in shooting right across the interspace of air in the direction imparted by the shove." –On the nature of the Universe

Despite being similar to later particle theories, Lucretius's views were not generally accepted.

Ptolemy (c. 2nd century) wrote about the refraction of light in his book Optics.^[15]

Physical theories

René Descartes (1596–1650) held that light was a mechanical property of the luminous body, rejecting the "forms" of Ibn al-Haytham and Witelo as well as the "species" of Bacon, Grosseteste, andKepler.^[16] In 1637 he published a theory of the refraction of light that assumed, incorrectly, that light travelled faster in a denser medium than in a less dense medium. Descartes arrived at this conclusion by analogy with the behaviour of sound waves.^{[citation needed]} Although Descartes was incorrect about the relative speeds, he was correct in assuming that light behaved like a wave and in concluding that refraction could be explained by the speed of light in different media.

Descartes is not the first to use the mechanical analogies but because he clearly asserts that light is only a mechanical property of the luminous body and the transmitting medium, Descartes' theory of light is regarded as the start of modern physical optics.^[17]

Particle theory

Main article: Corpuscular theory of light

Pierre Gassendi.

Pierre Gassendi (1592–1655), an atomist, proposed a particle theory of light which was published posthumously in the 1660s. Isaac Newton studied Gassendi's work at an early age, and preferred his view to Descartes' theory of the plenum. He stated in his Hypothesis of Light of 1675 that light was composed ofcorpuscles (particles of matter) which were emitted in all directions from a source. One of Newton's arguments against the wave nature of light was that waves were known to bend around obstacles, while light travelled only in straight lines. He did, however, explain the phenomenon of the diffraction of light (which had been observed by Francesco Grimaldi) by allowing that a light particle could create a localised wave in the aether.

Newton's theory could be used to predict the reflection of light, but could only explain refraction by incorrectly assuming that light accelerated upon entering a denser medium because the gravitational pull was greater. Newton published the final version of his theory in his Opticks of 1704. His reputation helped theparticle theory of light to hold sway during the 18th century. The particle theory of light led Laplace to argue that a body could be so massive that light could not escape from it. In other words it would become what is now called a black hole. Laplace withdrew his suggestion when the wave theory of light was firmly established. A translation of his essay appears in The large scale structure of space-time, by Stephen Hawking and George F. R. Ellis.

Wave theory

Main article: Electromagnetic wave

In the 1660s, Robert Hooke published a wave theory of light. Christiaan Huygens worked out his own wave theory of light in 1678, and published it in his Treatise on light in 1690. He proposed that light was emitted in all directions as a series of waves in a medium called the Luminiferous ether. As waves are not affected by gravity, it was assumed that they slowed down upon entering a denser medium.

Thomas Young's sketch of the two-slit experiment showing the diffractionof light. Young's experiments supported the theory that light consists of waves.

The wave theory predicted that light waves could interfere with each other like sound waves (as noted around 1800 by Thomas Young), and that light could bepolarised, if it were a transverse wave. Young showed by means of a diffraction experiment that light behaved as waves. He also proposed that different colourswere caused by different wavelengths of light, and explained colour vision in terms of three-coloured receptors in the eye.

Another supporter of the wave theory was Leonhard Euler. He argued in Nova theoria lucis et colorum (1746) that diffraction could more easily be explained by a wave theory.

Later, Augustin-Jean Fresnel independently worked out his own wave theory of light, and presented it to the Académie des Sciences in 1817. Simeon Denis Poisson added to Fresnel's mathematical work to produce a convincing argument in favour of the wave theory, helping to overturn Newton's corpuscular theory. By the year 1821, Fresnel was able to show via mathematical methods that polarisation could be explained only by the wave theory of light and only if light was entirely transverse, with no longitudinal vibration whatsoever.

The weakness of the wave theory was that light waves, like sound waves, would need a medium for transmission. A hypothetical substance called theluminiferous aether was proposed, but its existence was cast into strong doubt in the late nineteenth century by the Michelson-Morley experiment.

Newton's corpuscular theory implied that light would travel faster in a denser medium, while the wave theory of Huygens and others implied the opposite. At that time, the speed of light could not be measured accurately enough to decide which theory was correct. The first to make a sufficiently accurate measurement was Léon Foucault, in 1850.^[18] His result supported the wave theory, and the classical particle theory was finally abandoned.

Electromagnetic theory

Main article: Electromagnetism

A linearly polarised light wave frozen in time and showing the two oscillating components of light; an electric field and a magnetic fieldperpendicular to each other and to the direction of motion (a transverse wave).

In 1845, Michael Faraday discovered that the plane of polarisation of linearly polarised light is rotated when the light rays travel along the magnetic field direction in the presence of a transparent dielectric, an effect now known as Faraday rotation.^[19] This was the first evidence that light was related to electromagnetism. In 1846 he speculated that light might be some form of disturbance propagating along magnetic field lines.^[20] Faraday proposed in 1847 that light was a high-frequency electromagnetic vibration, which could propagate even in the absence of a medium such as the ether.

Faraday's work inspired James Clerk Maxwell to study electromagnetic radiation and light. Maxwell discovered that self-propagating electromagnetic waves would travel through space at a constant speed, which happened to be equal to the previously measured speed of light. From this, Maxwell concluded that light was a form of electromagnetic radiation: he first stated this result in 1862 inOn Physical Lines of Force. In 1873, he published A Treatise on Electricity and Magnetism, which contained a full mathematical description of the behaviour of electric and magnetic fields, still known as Maxwell's equations. Soon after, Heinrich Hertz confirmed Maxwell's theory experimentally by generating and detecting radio waves in the laboratory, and demonstrating that these waves behaved exactly like visible light, exhibiting properties such as reflection, refraction, diffraction, and interference. Maxwell's theory and Hertz's experiments led directly to the development of modern radio, radar, television, electromagnetic imaging, and wireless communications.

The special theory of relativity

Main article: Special theory of relativity

Albert Einstein had proposed the Theory of relativity.

The wave theory was successful in explaining nearly all optical and electromagnetic phenomena, and was a great triumph of nineteenth century physics. By the late nineteenth century, however, a handful of experimental anomalies remained that could not be explained by or were in direct conflict with the wave theory. One of these anomalies involved a controversy over the speed of light. The constant speed of light predicted by Maxwell's equations and confirmed by the Michelson-Morley experiment contradicted the mechanical laws of motion that had been unchallenged since the time of Galileo, which stated that all speeds were relative to the speed of the observer. In 1905, Albert Einstein resolved this paradox by proposing that space and time appeared to be changeable entities, which accounted for the constancy of the speed of light. Einstein also proposed a previously unknown fundamental equivalence between energy andmass with his famous equation

where E is energy, m is, depending on the context, the rest mass or the relativistic mass, and c is the speed of light in a vacuum.

Particle theory revisited

Another experimental anomaly was the photoelectric effect, by which light striking a metal surface ejected electrons from the surface, causing an electric current to flow across an applied voltage. Experimental measurements demonstrated that the energy of individual ejected electrons was proportional to thefrequency, rather than the intensity, of the light. Furthermore, below a certain minimum frequency, which depended on the particular metal, no current would flow regardless of the intensity. These observations appeared to contradict the wave theory, and for years physicists tried in vain to find an explanation. In 1905, Einstein solved this puzzle as well, this time by resurrecting the particle theory of light to explain the observed effect. Because of the preponderance of evidence in favor of the wave theory, however, Einstein's ideas were met initially with great skepticism among established physicists. But eventually Einstein's explanation of the photoelectric effect would triumph, and it ultimately formed the basis for wave–particle duality and much of quantum mechanics.

Quantum theory

A third anomaly that arose in the late 19th century involved a contradiction between the wave theory of light and measurements of the electromagnetic spectrum emitted by thermal radiators, or so-called black bodies. Physicists struggled with this problem, which later became known as the ultraviolet catastrophe, unsuccessfully for many years. In 1900, Max Planck developed a new theory ofblack-body radiation that explained the observed spectrum. Planck's theory was based on the idea that black bodies emit light (and other electromagnetic radiation) only as discrete bundles or packets of energy. These packets were called quanta, and the particle of light was given the name photon, to correspond with other particles being described around this time, such as the electron and proton. A photon has an energy, E, proportional to its frequency, f, by

where h is Planck's constant, λ is the wavelength and c is the speed of light. Likewise, the momentum p of a photon is also proportional to its frequency and inversely proportional to its wavelength:

As it originally stood, this theory did not explain the simultaneous wave- and particle-like natures of light, though Planck would later work on theories that did. In 1918, Planck received the Nobel Prize in Physics for his part in the founding of quantum theory.

Wave–particle duality

The modern theory that explains the nature of light includes the notion of wave–particle duality, described by Albert Einstein in the early 1900s, based on his study of the photoelectric effect and Planck's results. Einstein asserted that the energy of a photon is proportional to its frequency. More generally, the theory states that everything has both a particle nature and a wave nature, and various experiments can be done to bring out one or the other. The particle nature is more easily discerned if an object has a large mass, and it was not until a bold proposition by Louis de Broglie in 1924 that the scientific community realised that electrons also exhibited wave–particle duality. The wave nature of electrons was experimentally demonstrated by Davisson and Germer in 1927. Einstein received the Nobel Prize in 1921 for his work with the wave–particle duality on photons (especially explaining the photoelectric effect thereby), and de Broglie followed in 1929 for his extension to other particles.

Quantum electrodynamics

Main article: Quantum electrodynamics

The quantum mechanical theory of light and electromagnetic radiation continued to evolve through the 1920s and 1930s, and culminated with the development during the 1940s of the theory of quantum electrodynamics, or QED. This so-called quantum field theory is among the most comprehensive and experimentally successful theories ever formulated to explain a set of natural phenomena. QED was developed primarily by physicists Richard Feynman, Freeman Dyson, Julian Schwinger, and Shin-Ichiro Tomonaga. Feynman, Schwinger, and Tomonaga shared the 1965 Nobel Prize in Physics for their contributions.