Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes that are passed on from parent to offspring during reproduction. Different characteristics tend to exist within any given population as a result of mutation, genetic recombination and other sources of genetic variation. Evolution occurs when evolutionary processes such as natural selection and genetic drift act on this variation, resulting in certain characteristics becoming more common or rare within a population, it is this process of evolution that has given rise to biodiversity at every level of biological organisation, including the levels of species, individual organisms and molecules. The scientific theory of evolution by natural selection was proposed by Charles Darwin and Alfred Russel Wallace in the mid-19th century and was set out in detail in Darwin's book On the Origin of Species. Evolution by natural selection was first demonstrated by the observation that more offspring are produced than can survive.
This is followed by three observable facts about living organisms: 1) traits vary among individuals with respect to their morphology and behaviour, 2) different traits confer different rates of survival and reproduction and 3) traits can be passed from generation to generation. Thus, in successive generations members of a population are more to be replaced by the progenies of parents with favourable characteristics that have enabled them to survive and reproduce in their respective environments. In the early 20th century, other competing ideas of evolution such as mutationism and orthogenesis were refuted as the modern synthesis reconciled Darwinian evolution with classical genetics, which established adaptive evolution as being caused by natural selection acting on Mendelian genetic variation. All life on Earth shares a last universal common ancestor that lived 3.5–3.8 billion years ago. The fossil record includes a progression from early biogenic graphite, to microbial mat fossils, to fossilised multicellular organisms.
Existing patterns of biodiversity have been shaped by repeated formations of new species, changes within species and loss of species throughout the evolutionary history of life on Earth. Morphological and biochemical traits are more similar among species that share a more recent common ancestor, can be used to reconstruct phylogenetic trees. Evolutionary biologists have continued to study various aspects of evolution by forming and testing hypotheses as well as constructing theories based on evidence from the field or laboratory and on data generated by the methods of mathematical and theoretical biology, their discoveries have influenced not just the development of biology but numerous other scientific and industrial fields, including agriculture and computer science. The proposal that one type of organism could descend from another type goes back to some of the first pre-Socratic Greek philosophers, such as Anaximander and Empedocles; such proposals survived into Roman times. The poet and philosopher Lucretius followed Empedocles in his masterwork De rerum natura.
In contrast to these materialistic views, Aristotelianism considered all natural things as actualisations of fixed natural possibilities, known as forms. This was part of a medieval teleological understanding of nature in which all things have an intended role to play in a divine cosmic order. Variations of this idea became the standard understanding of the Middle Ages and were integrated into Christian learning, but Aristotle did not demand that real types of organisms always correspond one-for-one with exact metaphysical forms and gave examples of how new types of living things could come to be. In the 17th century, the new method of modern science rejected the Aristotelian approach, it sought explanations of natural phenomena in terms of physical laws that were the same for all visible things and that did not require the existence of any fixed natural categories or divine cosmic order. However, this new approach was slow to take root in the biological sciences, the last bastion of the concept of fixed natural types.
John Ray applied one of the more general terms for fixed natural types, "species," to plant and animal types, but he identified each type of living thing as a species and proposed that each species could be defined by the features that perpetuated themselves generation after generation. The biological classification introduced by Carl Linnaeus in 1735 explicitly recognised the hierarchical nature of species relationships, but still viewed species as fixed according to a divine plan. Other naturalists of this time speculated on the evolutionary change of species over time according to natural laws. In 1751, Pierre Louis Maupertuis wrote of natural modifications occurring during reproduction and accumulating over many generations to produce new species. Georges-Louis Leclerc, Comte de Buffon suggested that species could degenerate into different organisms, Erasmus Darwin proposed that all warm-blooded animals could have descended from a single microorganism; the first full-fledged evolutionary scheme was Jean-Baptiste Lamarck's "transmutation" theory of 1809, which envisaged spontaneous generation continually producing simple forms of life that developed greater complexity in parallel lineages with an inherent progressive tendency, postulated that on a local level, these lineages adapted to the environment by inheriting changes caused by their use or disuse in parents.
These ideas were cond
Cell division is the process by which a parent cell divides into two or more daughter cells. Cell division occurs as part of a larger cell cycle. In eukaryotes, there are two distinct types of cell division: a vegetative division, whereby each daughter cell is genetically identical to the parent cell, a reproductive cell division, whereby the number of chromosomes in the daughter cells is reduced by half to produce haploid gametes. Meiosis results in four haploid daughter cells by undergoing one round of DNA replication followed by two divisions. Homologous chromosomes are separated in the first division, sister chromatids are separated in the second division. Both of these cell division cycles are used in the process of sexual reproduction at some point in their life cycle. Both are believed to be present in the last eukaryotic common ancestor. Prokaryotes undergo a vegetative cell division known as binary fission, where their genetic material is segregated into two daughter cells. All cell divisions, regardless of organism, are preceded by a single round of DNA replication.
For simple unicellular microorganisms such as the amoeba, one cell division is equivalent to reproduction – an entire new organism is created. On a larger scale, mitotic cell division can create progeny from multicellular organisms, such as plants that grow from cuttings. Mitotic cell division enables sexually reproducing organisms to develop from the one-celled zygote, which itself was produced by meiotic cell division from gametes. After growth, cell division by mitosis allows for continual repair of the organism; the human body experiences about 10 quadrillion cell divisions in a lifetime. The primary concern of cell division is the maintenance of the original cell's genome. Before division can occur, the genomic information, stored in chromosomes must be replicated, the duplicated genome must be separated cleanly between cells. A great deal of cellular infrastructure is involved in keeping genomic information consistent between generations. Interphase is the process a cell must go through before mitosis and cytokinesis.
Interphase consists of three main stages: G1, S, G2. G1 is a time of growth for the cell where specialized cellular functions occur in order to prepare the cell for DNA Replication. There are checkpoints during interphase that allow the cell to be either progressed or denied further development. In S phase, the chromosomes are replicated in order for the genetic content to be maintained. During G2, the cell undergoes the final stages of growth before it enters the M phase, where spindles are synthesized; the M phase, can be either meiosis depending on the type of cell. Germ cells, or gametes, undergo meiosis. After the cell proceeds through the M phase, it may undergo cell division through cytokinesis; the control of each checkpoint is controlled by cyclin and cyclin dependent kinases. The progression of interphase is the result of the increased amount of cyclin; as the amount of cyclin increases and more cyclin dependent kinases attach to cyclin signaling the cell further into interphase. The peak of the cyclin attached to the cyclin dependent kinases this system pushes the cell out of interphase and into the M phase, where mitosis and cytokinesis occur.
There are three transition checkpoints. The most important being the G1-S transition checkpoint. If the cell does not pass this phase the cell will most not go through the rest of the cell division cycle. Prophase is the first stage of division; the nuclear envelope is broken down, long strands of chromatin condense to form shorter more visible strands called chromosomes, the nucleolus disappears, microtubules attach to the chromosomes at the kinetochores present in the centromere. Microtubules associated with the alignment and separation of chromosomes are referred to as the spindle and spindle fibers. Chromosomes will be visible under a microscope and will be connected at the centromere. During this condensation and alignment period, homologous over. In metaphase, the centromeres of the chromosomes convene themselves on the metaphase plate, an imaginary line, equidistant from the two centrosome poles. Chromosomes line up in the middle of the cell by MTOCs by pushing and pulling on centromeres of both chromatids which causes the chromosome to move to the center.
The chromosomes are still condensing and are at one step away from being the most coiled and condensed they will be. Spindle fibres have connected to the kinetochores. At this point, the chromosomes are ready to split into opposite poles of the cell towards the spindle to which they are connected. Anaphase is a short stage of the cell cycle and occurs after the chromosomes align at the mitotic plate. After the chromosomes line up in the middle of the cell, the spindle fibers will pull them apart; the chromosomes are split apart as the sister chromatids move to opposite sides of the cell. While the sister chromatids are being pulled apart and plasma gets elongated from non-kinetochore microtubules Telophase is the last stage of the cell cycle. A cleavage furrow splits the cell in two; these two cells form around the chromatin at the two poles of the cell. Two nuclear membranes begin to reform and the chromatin begin to unwind. Cells are broadly classified into two main categories: simple, non-nucleated prokaryotic cells, complex, nucleated eukaryotic cells.
Owing to their structural differences and prokaryotic cells do not divide in the same way. The pattern of cell division that tr
Phytochemistry is the study of phytochemicals, which are chemicals derived from plants. Those studying phytochemistry strive to describe the structures of the large number of secondary metabolic compounds found in plants, the functions of these compounds in human and plant biology, the biosynthesis of these compounds. Plants synthesize phytochemicals for many reasons, including to protect themselves against insect attacks and plant diseases. Phytochemicals in food plants are active in human biology, in many cases have health benefits; the compounds found in plants are of many kinds, but most are in four major biochemical classes, the alkaloids, glycosides and terpenes. Phytochemistry can be considered sub-fields of chemistry. Activities can be led in the wild with the aid of ethnobotany; the applications of the discipline can be for pharmacognosy, or the discovery of new drugs, or as an aid for plant physiology studies. Techniques used in the field of phytochemistry are extraction and structural elucidation of natural products, as well as various chromatography techniques.
The list of simple elements of which plants are constructed—carbon, hydrogen, phosphorus, etc.—is not different from similar lists for animals, fungi, or bacteria. The fundamental atomic components of plants are the same as for all life. Phytochemistry is used in the field of Chinese medicine in the field of herbal medicine. Phytochemical technique applies to the quality control of Chinese medicine, Ayurvedic medicine or herbal medicine of various chemical components, such as saponins, volatile oils and anthraquinones. In the development of rapid and reproducible analytical techniques, the combination of HPLC with different detectors, such as diode array detector, refractive index detector, evaporative light scattering detector and mass spectrometric detector, has been developed. In most cases, biologically active compounds in Chinese medicine, Ayurveda, or herbal medicine have not been determined. Therefore, it is important to use the phytochemical methods to screen and analyze bioactive components, not only for the quality control of crude drugs, but for the elucidation of their therapeutic mechanisms.
Modern pharmacological studies indicate that binding to receptors or ion channels on cell membranes is the first step of some drug actions. A new method in phytochemistry called biochromatography has been developed; this method combines human red cell membrane extraction and high performance liquid chromatography to screen potential active components in Chinese medicine. Many plants produce chemical compounds for defence against herbivores; these are useful as drugs, the content and known pharmacological activity of these substances in medicinal plants is the scientific basis for their use. The major classes of pharmacologically active phytochemicals are described below, with examples of medicinal plants that contain them. Human settlements are surrounded by weeds useful as medicines, such as nettle and chickweed. Many phytochemicals, including curcumin, epigallocatechin gallate and resveratrol are pan-assay interference compounds and are not useful in drug discovery. Alkaloids are bitter-tasting chemicals widespread in nature, toxic.
There are several classes with different modes of action as drugs, both recreational and pharmaceutical. Medicines of different classes include atropine and hyoscyamine, the traditional medicine berberine, cocaine, morphine, reserpine and quinine, vincristine. Anthraquinone glycosides are found in the laxatives senna and Aloe; the cardiac glycosides are powerful drugs from plants including lily of the valley. They include digoxin and digitoxin which support the beating of the heart, act as diuretics. Polyphenols of several classes are widespread in plants, they include the colourful anthocyanins, hormone-mimicking phytoestrogens, astringent tannins. In Ayurveda, the astringent rind of the pomegranate is used as a medicine, while polyphenol extracts from plant materials such as grape seeds are sold for their potential health benefits They have been continually studied in cell cultures for their different anti-cancer effects. Plants containing phytoestrogens have been used for centuries to treat gynaecological disorders such as fertility and menopausal problems.
Among these plants are Pueraria mirifica, angelica and anise. Terpenes and terpenoids of many kinds are found in resinous plants such as the conifers, they are aromatic and serve to repel herbivores. Their scent makes them useful in essential oils, whether for perfumes such as rose and lavender, or for aromatherapy; some have had medicinal uses: thymol is an antiseptic and was once used as a vermifuge. Tropical Botanical Garden and Research Institute UBC Botanical Garden and Centre for Plant Research
In biology and biochemistry, a lipid is a biomolecule, soluble in nonpolar solvents. Non-polar solvents are hydrocarbons used to dissolve other occurring hydrocarbon lipid molecules that do not dissolve in water, including fatty acids, sterols, fat-soluble vitamins, diglycerides and phospholipids; the functions of lipids include storing energy and acting as structural components of cell membranes. Lipids have applications in the food industries as well as in nanotechnology. Scientists sometimes broadly define lipids as amphiphilic small molecules. Biological lipids originate or in part from two distinct types of biochemical subunits or "building-blocks": ketoacyl and isoprene groups. Using this approach, lipids may be divided into eight categories: fatty acids, glycerophospholipids, sphingolipids and polyketides. Although the term "lipid" is sometimes used as a synonym for fats, fats are a subgroup of lipids called triglycerides. Lipids encompass molecules such as fatty acids and their derivatives, as well as other sterol-containing metabolites such as cholesterol.
Although humans and other mammals use various biosynthetic pathways both to break down and to synthesize lipids, some essential lipids can't be made this way and must be obtained from the diet. In 1815, Henry Braconnot classified lipids in two categories and huiles. In 1823, Michel Eugène Chevreul developed a more detailed classification, including oils, tallow, resins and volatile oils. In 1827, William Prout recognized fat, along with protein and carbohydrate, as an important nutrient for humans and animals. For a century, chemists regarded "fats" as only simple lipids made of fatty acids and glycerol, but new forms were described later. Theodore Gobley discovered phospholipids in mammalian brain and hen egg, called by him as "lecithins". Thudichum discovered in human brain some phospholipids and sphingolipids; the terms lipoid, lipin and lipid have been used with varied meanings from author to author. In 1912, Rosenbloom and Gies proposed the substitution of "lipoid" by "lipin". In 1920, Bloor introduced a new classification for "lipoids": simple lipoids, compound lipoids, the derived lipoids.
The word "lipid", which stems etymologically from the Greek lipos, was introduced in 1923 by Gabriel Bertrand. Bertrands included in the concept not only the traditional fats, but the "lipoids", with a complex constitution. In 1947, T. P. Hilditch divided lipids into "simple lipids", with greases and waxes, "complex lipids", with phospholipids and glycolipids. Fatty acids, or fatty acid residues when they are part of a lipid, are a diverse group of molecules synthesized by chain-elongation of an acetyl-CoA primer with malonyl-CoA or methylmalonyl-CoA groups in a process called fatty acid synthesis, they are made of a hydrocarbon chain. The fatty acid structure is one of the most fundamental categories of biological lipids, is used as a building-block of more structurally complex lipids; the carbon chain between four and 24 carbons long, may be saturated or unsaturated, may be attached to functional groups containing oxygen, halogens and sulfur. If a fatty acid contains a double bond, there is the possibility of either a cis or trans geometric isomerism, which affects the molecule's configuration.
Cis-double bonds cause the fatty acid chain to bend, an effect, compounded with more double bonds in the chain. Three double bonds in 18-carbon linolenic acid, the most abundant fatty-acyl chains of plant thylakoid membranes, render these membranes fluid despite environmental low-temperatures, makes linolenic acid give dominating sharp peaks in high resolution 13-C NMR spectra of chloroplasts; this in turn plays an important role in the function of cell membranes. Most occurring fatty acids are of the cis configuration, although the trans form does exist in some natural and hydrogenated fats and oils. Examples of biologically important fatty acids include the eicosanoids, derived from arachidonic acid and eicosapentaenoic acid, that include prostaglandins and thromboxanes. Docosahexaenoic acid is important in biological systems with respect to sight. Other major lipid classes in the fatty acid category are the fatty esters and fatty amides. Fatty esters include important biochemical intermediates such as wax esters, fatty acid thioester coenzyme A derivatives, fatty acid thioester ACP derivatives and fatty acid carnitines.
The fatty amides include N-acyl ethanolamines, such as the cannabinoid neurotransmitter anandamide. Glycerolipids are composed of mono-, di-, tri-substituted glycerols, the best-known being the fatty acid triesters of glycerol, called triglycerides; the word "triacylgl
A macromolecule is a large molecule, such as protein created by the polymerization of smaller subunits. They are composed of thousands of atoms or more; the most common macromolecules in biochemistry are large non-polymeric molecules. Synthetic macromolecules include common plastics and synthetic fibers as well as experimental materials such as carbon nanotubes; the term macromolecule was coined by Nobel laureate Hermann Staudinger in the 1920s, although his first relevant publication on this field only mentions high molecular compounds. At that time the phrase polymer, as introduced by Berzelius in 1833, had a different meaning from that of today: it was another form of isomerism for example with benzene and acetylene and had little to do with size. Usage of the term to describe large molecules varies among the disciplines. For example, while biology refers to macromolecules as the four large molecules comprising living things, in chemistry, the term may refer to aggregates of two or more molecules held together by intermolecular forces rather than covalent bonds but which do not dissociate.
According to the standard IUPAC definition, the term macromolecule as used in polymer science refers only to a single molecule. For example, a single polymeric molecule is appropriately described as a "macromolecule" or "polymer molecule" rather than a "polymer," which suggests a substance composed of macromolecules; because of their size, macromolecules are not conveniently described in terms of stoichiometry alone. The structure of simple macromolecules, such as homopolymers, may be described in terms of the individual monomer subunit and total molecular mass. Complicated biomacromolecules, on the other hand, require multi-faceted structural description such as the hierarchy of structures used to describe proteins. In British English, the word "macromolecule" tends to be called "high polymer". Macromolecules have unusual physical properties that do not occur for smaller molecules. Another common macromolecular property that does not characterize smaller molecules is their relative insolubility in water and similar solvents, instead forming colloids.
Many require particular ions to dissolve in water. Many proteins will denature if the solute concentration of their solution is too high or too low. High concentrations of macromolecules in a solution can alter the rates and equilibrium constants of the reactions of other macromolecules, through an effect known as macromolecular crowding; this comes from macromolecules excluding other molecules from a large part of the volume of the solution, thereby increasing the effective concentrations of these molecules. All living organisms are dependent on three essential biopolymers for their biological functions: DNA, RNA and proteins; each of these molecules is required for life since each plays a distinct, indispensable role in the cell. The simple summary is that DNA makes RNA, RNA makes proteins. DNA, RNA, proteins all consist of a repeating structure of related building blocks. In general, they are all unbranched polymers, so can be represented in the form of a string. Indeed, they can be viewed as a string of beads, with each bead representing a single nucleotide or amino acid monomer linked together through covalent chemical bonds into a long chain.
In most cases, the monomers within the chain have a strong propensity to interact with other amino acids or nucleotides. In DNA and RNA, this can take the form of Watson-Crick base pairs, although many more complicated interactions can and do occur; because of the double-stranded nature of DNA all of the nucleotides take the form of Watson-Crick base pairs between nucleotides on the two complementary strands of the double-helix. In contrast, both RNA and proteins are single-stranded. Therefore, they are not constrained by the regular geometry of the DNA double helix, so fold into complex three-dimensional shapes dependent on their sequence; these different shapes are responsible for many of the common properties of RNA and proteins, including the formation of specific binding pockets, the ability to catalyse biochemical reactions. DNA is an information storage macromolecule that encodes the complete set of instructions that are required to assemble and reproduce every living organism. DNA and RNA are both capable of encoding genetic information, because there are biochemical mechanisms which read the information coded within a DNA or RNA sequence and use it to generate a specified protein.
On the other hand, the sequence information of a protein molecule is not used by cells to functionally encode genetic information. DNA has three primary attributes that allow it to be far better than RNA at encoding genetic information. First, it is double-stranded, so that there are a minimum of two copies of the information encoding each gene in every cell. Second, DNA has a much greater stability against breakdown than does RNA, an attribute associated with the absence of the 2'-hydroxyl group within every nucleotide of DNA. Third sophisticated DNA surveillance and repair systems are present which monitor damage to the DNA and repair the sequence when necessary. Analogous systems have not evolved for repairing damaged RNA molecules. Chromosomes can contain many billions of atoms, arranged in a specific chemical structure. Proteins are functional macromolecules responsible for catalysing the biochemical reactions that sustain life. Proteins carry out all functions of an organism, for example p
Molecular biology is a branch of biology that concerns the molecular basis of biological activity between biomolecules in the various systems of a cell, including the interactions between DNA, RNA, proteins and their biosynthesis, as well as the regulation of these interactions. Writing in Nature in 1961, William Astbury described molecular biology as:...not so much a technique as an approach, an approach from the viewpoint of the so-called basic sciences with the leading idea of searching below the large-scale manifestations of classical biology for the corresponding molecular plan. It is concerned with the forms of biological molecules and is predominantly three-dimensional and structural – which does not mean, that it is a refinement of morphology, it must at the same time inquire into function. Researchers in molecular biology use specific techniques native to molecular biology but combine these with techniques and ideas from genetics and biochemistry. There is not a defined line between these disciplines.
This is shown in the following schematic that depicts one possible view of the relationships between the fields: Biochemistry is the study of the chemical substances and vital processes occurring in live organisms. Biochemists focus on the role and structure of biomolecules; the study of the chemistry behind biological processes and the synthesis of biologically active molecules are examples of biochemistry. Genetics is the study of the effect of genetic differences in organisms; this can be inferred by the absence of a normal component. The study of "mutants" – organisms which lack one or more functional components with respect to the so-called "wild type" or normal phenotype. Genetic interactions can confound simple interpretations of such "knockout" studies. Molecular biology is the study of molecular underpinnings of the processes of replication, transcription and cell function; the central dogma of molecular biology where genetic material is transcribed into RNA and translated into protein, despite being oversimplified, still provides a good starting point for understanding the field.
The picture has been revised in light of emerging novel roles for RNA. Much of molecular biology is quantitative, much work has been done at its interface with computer science in bioinformatics and computational biology. In the early 2000s, the study of gene structure and function, molecular genetics, has been among the most prominent sub-fields of molecular biology. Many other areas of biology focus on molecules, either directly studying interactions in their own right such as in cell biology and developmental biology, or indirectly, where molecular techniques are used to infer historical attributes of populations or species, as in fields in evolutionary biology such as population genetics and phylogenetics. There is a long tradition of studying biomolecules "from the ground up" in biophysics. One of the most basic techniques of molecular biology to study protein function is molecular cloning. In this technique, DNA coding for a protein of interest is cloned using polymerase chain reaction, and/or restriction enzymes into a plasmid.
A vector has 3 distinctive features: an origin of replication, a multiple cloning site, a selective marker antibiotic resistance. Located upstream of the multiple cloning site are the promoter regions and the transcription start site which regulate the expression of cloned gene; this plasmid can be inserted into either bacterial or animal cells. Introducing DNA into bacterial cells can be done by transformation via uptake of naked DNA, conjugation via cell-cell contact or by transduction via viral vector. Introducing DNA into eukaryotic cells, such as animal cells, by physical or chemical means is called transfection. Several different transfection techniques are available, such as calcium phosphate transfection, electroporation and liposome transfection; the plasmid may be integrated into the genome, resulting in a stable transfection, or may remain independent of the genome, called transient transfection. DNA coding for a protein of interest is now inside a cell, the protein can now be expressed.
A variety of systems, such as inducible promoters and specific cell-signaling factors, are available to help express the protein of interest at high levels. Large quantities of a protein can be extracted from the bacterial or eukaryotic cell; the protein can be tested for enzymatic activity under a variety of situations, the protein may be crystallized so its tertiary structure can be studied, or, in the pharmaceutical industry, the activity of new drugs against the protein can be studied. Polymerase chain reaction is an versatile technique for copying DNA. In brief, PCR allows a specific DNA sequence to be modified in predetermined ways; the reaction is powerful and under perfect conditions could amplify one DNA molecule to become 1.07 billion molecules in less than two hours. The PCR technique can be used to introduce restriction enzyme sites to ends of DNA molecules, or to mutate particular bases of DNA, the latter is a method referred to as site-directed mutagenesis. PCR can be used to determine whether a particular DNA fragment is found in a cDNA library.
PCR has many variations, like reverse transcription PCR for amplification of RNA, more quantitative PCR which allow for quantitative measurement of DNA or RNA molecules. Gel electrophoresis is one of the principal tools of molecular biology; the basic principle is that DNA, RNA, proteins can all be separated by means of an electric field and size. In agarose gel electrophoresis, DNA and RNA can be separated on th
Carbon is a chemical element with symbol C and atomic number 6. It is nonmetallic and tetravalent—making four electrons available to form covalent chemical bonds, it belongs to group 14 of the periodic table. Three isotopes occur 12C and 13C being stable, while 14C is a radionuclide, decaying with a half-life of about 5,730 years. Carbon is one of the few elements known since antiquity. Carbon is the 15th most abundant element in the Earth's crust, the fourth most abundant element in the universe by mass after hydrogen and oxygen. Carbon's abundance, its unique diversity of organic compounds, its unusual ability to form polymers at the temperatures encountered on Earth enables this element to serve as a common element of all known life, it is the second most abundant element in the human body by mass after oxygen. The atoms of carbon can bond together in different ways, termed allotropes of carbon; the best known are graphite and amorphous carbon. The physical properties of carbon vary with the allotropic form.
For example, graphite is opaque and black while diamond is transparent. Graphite is soft enough to form a streak on paper, while diamond is the hardest occurring material known. Graphite is a good electrical conductor. Under normal conditions, carbon nanotubes, graphene have the highest thermal conductivities of all known materials. All carbon allotropes are solids under normal conditions, with graphite being the most thermodynamically stable form at standard temperature and pressure, they are chemically resistant and require high temperature to react with oxygen. The most common oxidation state of carbon in inorganic compounds is +4, while +2 is found in carbon monoxide and transition metal carbonyl complexes; the largest sources of inorganic carbon are limestones and carbon dioxide, but significant quantities occur in organic deposits of coal, peat and methane clathrates. Carbon forms a vast number of compounds, more than any other element, with ten million compounds described to date, yet that number is but a fraction of the number of theoretically possible compounds under standard conditions.
For this reason, carbon has been referred to as the "king of the elements". The allotropes of carbon include graphite, one of the softest known substances, diamond, the hardest occurring substance, it bonds with other small atoms, including other carbon atoms, is capable of forming multiple stable covalent bonds with suitable multivalent atoms. Carbon is known to form ten million different compounds, a large majority of all chemical compounds. Carbon has the highest sublimation point of all elements. At atmospheric pressure it has no melting point, as its triple point is at 10.8±0.2 MPa and 4,600 ± 300 K, so it sublimes at about 3,900 K. Graphite is much more reactive than diamond at standard conditions, despite being more thermodynamically stable, as its delocalised pi system is much more vulnerable to attack. For example, graphite can be oxidised by hot concentrated nitric acid at standard conditions to mellitic acid, C66, which preserves the hexagonal units of graphite while breaking up the larger structure.
Carbon sublimes in a carbon arc, which has a temperature of about 5800 K. Thus, irrespective of its allotropic form, carbon remains solid at higher temperatures than the highest-melting-point metals such as tungsten or rhenium. Although thermodynamically prone to oxidation, carbon resists oxidation more than elements such as iron and copper, which are weaker reducing agents at room temperature. Carbon is the sixth element, with a ground-state electron configuration of 1s22s22p2, of which the four outer electrons are valence electrons, its first four ionisation energies, 1086.5, 2352.6, 4620.5 and 6222.7 kJ/mol, are much higher than those of the heavier group-14 elements. The electronegativity of carbon is 2.5 higher than the heavier group-14 elements, but close to most of the nearby nonmetals, as well as some of the second- and third-row transition metals. Carbon's covalent radii are taken as 77.2 pm, 66.7 pm and 60.3 pm, although these may vary depending on coordination number and what the carbon is bonded to.
In general, covalent radius decreases with higher bond order. Carbon compounds form the basis of all known life on Earth, the carbon–nitrogen cycle provides some of the energy produced by the Sun and other stars. Although it forms an extraordinary variety of compounds, most forms of carbon are comparatively unreactive under normal conditions. At standard temperature and pressure, it resists all but the strongest oxidizers, it does not react with hydrochloric acid, chlorine or any alkalis. At elevated temperatures, carbon reacts with oxygen to form carbon oxides and will rob oxygen from metal oxides to leave the elemental metal; this exothermic reaction is used in the iron and steel industry to smelt iron and to control the carbon content of steel: Fe3O4 + 4 C → 3 Fe + 4 COCarbon monoxide can be recycled to smelt more iron: Fe3O4 + 4 CO → 3 Fe + 4 CO2with sulfur to form carbon disulfide and with steam in the coal-gas reaction: C + H2O → CO + H2. Carbon combines with some metals at high temperatures to form metallic carbides, such as the iron carbide cementite in steel and tungsten carbide used as an abrasive and for making hard tips for cutting tools.
The system of carbon allotropes spans a range of extremes: Atomic carbon is a ver