174-420: See § Phyla Bacteria ( / b æ k ˈ t ɪər i ə / ; sg. : bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell . They constitute a large domain of prokaryotic microorganisms . Typically a few micrometres in length, bacteria were among the first life forms to appear on Earth , and are present in most of its habitats . Bacteria inhabit
348-411: A Honey-comb , but that the pores of it were not regular". To further support his theory, Matthias Schleiden and Theodor Schwann both also studied cells of both animal and plants. What they discovered were significant differences between the two types of cells. This put forth the idea that cells were not only fundamental to plants, but animals as well. Evolution Evolution is the change in
522-420: A membrane that envelops the cell, regulates what moves in and out (selectively permeable), and maintains the electric potential of the cell . Inside the membrane, the cytoplasm takes up most of the cell's volume. Except red blood cells , which lack a cell nucleus and most organelles to accommodate maximum space for hemoglobin , all cells possess DNA , the hereditary material of genes , and RNA , containing
696-641: A nucleus , and prokaryotic cells , which lack a nucleus but have a nucleoid region. Prokaryotes are single-celled organisms such as bacteria , whereas eukaryotes can be either single-celled, such as amoebae , or multicellular , such as some algae , plants , animals , and fungi . Eukaryotic cells contain organelles including mitochondria , which provide energy for cell functions; chloroplasts , which create sugars by photosynthesis , in plants; and ribosomes , which synthesise proteins. Cells were discovered by Robert Hooke in 1665, who named them after their resemblance to cells inhabited by Christian monks in
870-516: A nucleus , and other membrane-bound organelles . The DNA of a prokaryotic cell consists of a single circular chromosome that is in direct contact with the cytoplasm . The nuclear region in the cytoplasm is called the nucleoid . Most prokaryotes are the smallest of all organisms, ranging from 0.5 to 2.0 μm in diameter. A prokaryotic cell has three regions: Plants , animals , fungi , slime moulds , protozoa , and algae are all eukaryotic . These cells are about fifteen times wider than
1044-540: A potential difference analogous to a battery. The general lack of internal membranes in bacteria means these reactions, such as electron transport , occur across the cell membrane between the cytoplasm and the outside of the cell or periplasm . However, in many photosynthetic bacteria, the plasma membrane is highly folded and fills most of the cell with layers of light-gathering membrane. These light-gathering complexes may even form lipid-enclosed structures called chlorosomes in green sulfur bacteria . Bacteria do not have
1218-403: A terminal electron acceptor in a redox reaction . Chemotrophs are further divided by the types of compounds they use to transfer electrons. Bacteria that derive electrons from inorganic compounds such as hydrogen, carbon monoxide , or ammonia are called lithotrophs , while those that use organic compounds are called organotrophs . Still, more specifically, aerobic organisms use oxygen as
1392-490: A DNA sequence within a chromosome is known as a locus . If the DNA sequence at a locus varies between individuals, the different forms of this sequence are called alleles. DNA sequences can change through mutations, producing new alleles. If a mutation occurs within a gene, the new allele may affect the trait that the gene controls, altering the phenotype of the organism. However, while this simple correspondence between an allele and
1566-570: A GC-biased E. coli mutator strain in 1967, along with the proposal of the neutral theory , established the plausibility of mutational explanations for molecular patterns, which are now common in the molecular evolution literature. For instance, mutation biases are frequently invoked in models of codon usage. Such models also include effects of selection, following the mutation-selection-drift model, which allows both for mutation biases and differential selection based on effects on translation. Hypotheses of mutation bias have played an important role in
1740-742: A bacterial strain. However, liquid growth media are used when the measurement of growth or large volumes of cells are required. Growth in stirred liquid media occurs as an even cell suspension, making the cultures easy to divide and transfer, although isolating single bacteria from liquid media is difficult. The use of selective media (media with specific nutrients added or deficient, or with antibiotics added) can help identify specific organisms. Most laboratory techniques for growing bacteria use high levels of nutrients to produce large amounts of cells cheaply and quickly. However, in natural environments, nutrients are limited, meaning that bacteria cannot continue to reproduce indefinitely. This nutrient limitation has led
1914-454: A cell. Some (such as the nucleus and Golgi apparatus ) are typically solitary, while others (such as mitochondria , chloroplasts , peroxisomes and lysosomes ) can be numerous (hundreds to thousands). The cytosol is the gelatinous fluid that fills the cell and surrounds the organelles. Many cells also have structures which exist wholly or partially outside the cell membrane. These structures are notable because they are not protected from
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#17328023256752088-514: A chromosome becoming duplicated (usually by genetic recombination ), which can introduce extra copies of a gene into a genome. Extra copies of genes are a major source of the raw material needed for new genes to evolve. This is important because most new genes evolve within gene families from pre-existing genes that share common ancestors. For example, the human eye uses four genes to make structures that sense light: three for colour vision and one for night vision ; all four are descended from
2262-434: A discrete nucleus, usually with additional genetic material in some organelles like mitochondria and chloroplasts (see endosymbiotic theory ). A human cell has genetic material contained in the cell nucleus (the nuclear genome ) and in the mitochondria (the mitochondrial genome ). In humans, the nuclear genome is divided into 46 linear DNA molecules called chromosomes , including 22 homologous chromosome pairs and
2436-422: A disorganised slime layer of extracellular polymeric substances to a highly structured capsule . These structures can protect cells from engulfment by eukaryotic cells such as macrophages (part of the human immune system ). They can also act as antigens and be involved in cell recognition, as well as aiding attachment to surfaces and the formation of biofilms. The assembly of these extracellular structures
2610-407: A diverse range of single-celled organisms. The plants were created around 1.6 billion years ago with a second episode of symbiogenesis that added chloroplasts , derived from cyanobacteria . In 1665, Robert Hooke examined a thin slice of cork under his microscope , and saw a structure of small enclosures. He wrote "I could exceeding plainly perceive it to be all perforated and porous, much like
2784-506: A few hundred to a few thousand genes. The genes in bacterial genomes are usually a single continuous stretch of DNA. Although several different types of introns do exist in bacteria, these are much rarer than in eukaryotes. Bacteria, as asexual organisms, inherit an identical copy of the parent's genome and are clonal . However, all bacteria can evolve by selection on changes to their genetic material DNA caused by genetic recombination or mutations . Mutations arise from errors made during
2958-411: A few micrometres in thickness to up to half a metre in depth, and may contain multiple species of bacteria, protists and archaea. Bacteria living in biofilms display a complex arrangement of cells and extracellular components, forming secondary structures, such as microcolonies , through which there are networks of channels to enable better diffusion of nutrients. In natural environments, such as soil or
3132-411: A few species are visible to the unaided eye—for example, Thiomargarita namibiensis is up to half a millimetre long, Epulopiscium fishelsoni reaches 0.7 mm, and Thiomargarita magnifica can reach even 2 cm in length, which is 50 times larger than other known bacteria. Among the smallest bacteria are members of the genus Mycoplasma , which measure only 0.3 micrometres, as small as
3306-424: A fixed size and then reproduce through binary fission , a form of asexual reproduction . Under optimal conditions, bacteria can grow and divide extremely rapidly, and some bacterial populations can double as quickly as every 17 minutes. In cell division, two identical clone daughter cells are produced. Some bacteria, while still reproducing asexually, form more complex reproductive structures that help disperse
3480-414: A fluid mosaic membrane. Embedded within this membrane is a macromolecular structure called the porosome the universal secretory portal in cells and a variety of protein molecules that act as channels and pumps that move different molecules into and out of the cell. The membrane is semi-permeable, and selectively permeable, in that it can either let a substance ( molecule or ion ) pass through freely, to
3654-405: A functional three-dimensional protein molecule. Unicellular organisms can move in order to find food or escape predators. Common mechanisms of motion include flagella and cilia . In multicellular organisms, cells can move during processes such as wound healing, the immune response and cancer metastasis . For example, in wound healing in animals, white blood cells move to the wound site to kill
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#17328023256753828-400: A growing problem. Bacteria are important in sewage treatment and the breakdown of oil spills , the production of cheese and yogurt through fermentation , the recovery of gold, palladium , copper and other metals in the mining sector ( biomining , bioleaching ), as well as in biotechnology , and the manufacture of antibiotics and other chemicals. Once regarded as plants constituting
4002-427: A limited extent or not at all. Cell surface membranes also contain receptor proteins that allow cells to detect external signaling molecules such as hormones . 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
4176-402: A manner similar to a sequence of letters spelling out a sentence. Before a cell divides, the DNA is copied, so that each of the resulting two cells will inherit the DNA sequence. Portions of a DNA molecule that specify a single functional unit are called genes; different genes have different sequences of bases. Within cells, each long strand of DNA is called a chromosome . The specific location of
4350-407: A membrane-bound nucleus, and their genetic material is typically a single circular bacterial chromosome of DNA located in the cytoplasm in an irregularly shaped body called the nucleoid . The nucleoid contains the chromosome with its associated proteins and RNA . Like all other organisms , bacteria contain ribosomes for the production of proteins, but the structure of the bacterial ribosome
4524-427: A monastery. Cell theory , developed in 1839 by Matthias Jakob Schleiden and Theodor Schwann , states that all organisms are composed of one or more cells, that cells are the fundamental unit of structure and function in all living organisms, and that all cells come from pre-existing cells. Cells are broadly categorized into two types: eukaryotic cells , which possess a nucleus , and prokaryotic cells , which lack
4698-435: A nucleus but have a nucleoid region. Prokaryotes are single-celled organisms , whereas eukaryotes can be either single-celled or multicellular . Prokaryotes include bacteria and archaea , two of the three domains of life . Prokaryotic cells were the first form of life on Earth, characterized by having vital biological processes including cell signaling . They are simpler and smaller than eukaryotic cells, and lack
4872-405: A nutrient in a long-term laboratory experiment , Flavobacterium evolving a novel enzyme that allows these bacteria to grow on the by-products of nylon manufacturing, and the soil bacterium Sphingobium evolving an entirely new metabolic pathway that degrades the synthetic pesticide pentachlorophenol . An interesting but still controversial idea is that some adaptations might increase
5046-436: A pair of sex chromosomes . The mitochondrial genome is a circular DNA molecule distinct from nuclear DNA. Although the mitochondrial DNA is very small compared to nuclear chromosomes, 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
5220-525: A particular organism or group of organisms ( syntrophy ). Bacterial growth follows four phases. When a population of bacteria first enter a high-nutrient environment that allows growth, the cells need to adapt to their new environment. The first phase of growth is the lag phase , a period of slow growth when the cells are adapting to the high-nutrient environment and preparing for fast growth. The lag phase has high biosynthesis rates, as proteins necessary for rapid growth are produced. The second phase of growth
5394-611: A phenomenon termed de novo gene birth . The generation of new genes can also involve small parts of several genes being duplicated, with these fragments then recombining to form new combinations with new functions ( exon shuffling ). When new genes are assembled from shuffling pre-existing parts, domains act as modules with simple independent functions, which can be mixed together to produce new combinations with new and complex functions. For example, polyketide synthases are large enzymes that make antibiotics ; they contain up to 100 independent domains that each catalyse one step in
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5568-538: A population are therefore more likely to be replaced by the offspring of parents with favourable characteristics for that environment. In the early 20th century, competing ideas of evolution were refuted and evolution was combined with Mendelian inheritance and population genetics to give rise to modern evolutionary theory. In this synthesis the basis for heredity is in DNA molecules that pass information from generation to generation. The processes that change DNA in
5742-575: A population include natural selection, genetic drift, mutation , and gene flow . All life on Earth—including humanity —shares a last universal common ancestor (LUCA), which lived approximately 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 ( speciation ), changes within species ( anagenesis ), and loss of species ( extinction ) throughout
5916-430: A population is not critical, but instead a measure known as the effective population size. The effective population is usually smaller than the total population since it takes into account factors such as the level of inbreeding and the stage of the lifecycle in which the population is the smallest. The effective population size may not be the same for every gene in the same population. It is usually difficult to measure
6090-433: A population through neutral transitions due to the principles of excess capacity, presuppression, and ratcheting, and it has been applied in areas ranging from the origins of the spliceosome to the complex interdependence of microbial communities . The time it takes a neutral allele to become fixed by genetic drift depends on population size; fixation is more rapid in smaller populations. The number of individuals in
6264-461: A process called eukaryogenesis . This is widely agreed to have involved symbiogenesis , in which archaea and bacteria came together to create the first eukaryotic common ancestor. This cell had a new level of complexity and capability, with a nucleus and facultatively aerobic mitochondria . It evolved some 2 billion years ago into a population of single-celled organisms that included the last eukaryotic common ancestor, gaining capabilities along
6438-412: A process called transformation . Many bacteria can naturally take up DNA from the environment, while others must be chemically altered in order to induce them to take up DNA. The development of competence in nature is usually associated with stressful environmental conditions and seems to be an adaptation for facilitating repair of DNA damage in recipient cells. Second, bacteriophages can integrate into
6612-443: 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, always happens when a cell divides through mitosis or binary fission. This occurs during
6786-499: A range of genes from bacteria, fungi and plants. Viruses can also carry DNA between organisms, allowing transfer of genes even across biological domains . Large-scale gene transfer has also occurred between the ancestors of eukaryotic cells and bacteria, during the acquisition of chloroplasts and mitochondria . It is possible that eukaryotes themselves originated from horizontal gene transfers between bacteria and archaea . Some heritable changes cannot be explained by changes to
6960-430: A reduction in scope when it was discovered that (1) GC-biased gene conversion makes an important contribution to composition in diploid organisms such as mammals and (2) bacterial genomes frequently have AT-biased mutation. Contemporary thinking about the role of mutation biases reflects a different theory from that of Haldane and Fisher. More recent work showed that the original "pressures" theory assumes that evolution
7134-452: A single ancestral gene. New genes can be generated from an ancestral gene when a duplicate copy mutates and acquires a new function. This process is easier once a gene has been duplicated because it increases the redundancy of the system; one gene in the pair can acquire a new function while the other copy continues to perform its original function. Other types of mutations can even generate entirely new genes from previously noncoding DNA,
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7308-419: A single chromosome compared to expectations , which is called their linkage disequilibrium . A set of alleles that is usually inherited in a group is called a haplotype . This can be important when one allele in a particular haplotype is strongly beneficial: natural selection can drive a selective sweep that will also cause the other alleles in the haplotype to become more common in the population; this effect
7482-676: A single circular chromosome that can range in size from only 160,000 base pairs in the endosymbiotic bacteria Carsonella ruddii , to 12,200,000 base pairs (12.2 Mbp) in the soil-dwelling bacteria Sorangium cellulosum . There are many exceptions to this; for example, some Streptomyces and Borrelia species contain a single linear chromosome, while some Vibrio species contain more than one chromosome. Some bacteria contain plasmids , small extra-chromosomal molecules of DNA that may contain genes for various useful functions such as antibiotic resistance , metabolic capabilities, or various virulence factors . Bacteria genomes usually encode
7656-645: A source of electrons and a substrate for carbon anabolism . In many ways, bacterial metabolism provides traits that are useful for ecological stability and for human society. For example, diazotrophs have the ability to fix nitrogen gas using the enzyme nitrogenase . This trait, which can be found in bacteria of most metabolic types listed above, leads to the ecologically important processes of denitrification , sulfate reduction , and acetogenesis , respectively. Bacterial metabolic processes are important drivers in biological responses to pollution ; for example, sulfate-reducing bacteria are largely responsible for
7830-420: A species or population, in particular shifts in allele frequency and adaptation. Macroevolution is the outcome of long periods of microevolution. Thus, the distinction between micro- and macroevolution is not a fundamental one—the difference is simply the time involved. However, in macroevolution, the traits of the entire species may be important. For instance, a large amount of variation among individuals allows
8004-544: A species to rapidly adapt to new habitats , lessening the chance of it going extinct, while a wide geographic range increases the chance of speciation, by making it more likely that part of the population will become isolated. In this sense, microevolution and macroevolution might involve selection at different levels—with microevolution acting on genes and organisms, versus macroevolutionary processes such as species selection acting on entire species and affecting their rates of speciation and extinction. A common misconception
8178-411: A striking example are people with the inherited trait of albinism , who do not tan at all and are very sensitive to sunburn . Heritable characteristics are passed from one generation to the next via DNA , a molecule that encodes genetic information. DNA is a long biopolymer composed of four types of bases. The sequence of bases along a particular DNA molecule specifies the genetic information, in
8352-472: A system in which organisms interact with every other element, physical as well as biological , in their local environment. Eugene Odum , a founder of ecology, defined an ecosystem as: "Any unit that includes all of the organisms...in a given area interacting with the physical environment so that a flow of energy leads to clearly defined trophic structure, biotic diversity, and material cycles (i.e., exchange of materials between living and nonliving parts) within
8526-793: A thick peptidoglycan cell wall like a Gram-positive bacterium, but also a second outer layer of lipids. In many bacteria, an S-layer of rigidly arrayed protein molecules covers the outside of the cell. This layer provides chemical and physical protection for the cell surface and can act as a macromolecular diffusion barrier . S-layers have diverse functions and are known to act as virulence factors in Campylobacter species and contain surface enzymes in Bacillus stearothermophilus . Flagella are rigid protein structures, about 20 nanometres in diameter and up to 20 micrometres in length, that are used for motility . Flagella are driven by
8700-416: A three- dimensional random walk . Bacterial species differ in the number and arrangement of flagella on their surface; some have a single flagellum ( monotrichous ), a flagellum at each end ( amphitrichous ), clusters of flagella at the poles of the cell ( lophotrichous ), while others have flagella distributed over the entire surface of the cell ( peritrichous ). The flagella of a group of bacteria,
8874-428: A trait works in some cases, most traits are influenced by multiple genes in a quantitative or epistatic manner. Evolution can occur if there is genetic variation within a population. Variation comes from mutations in the genome, reshuffling of genes through sexual reproduction and migration between populations ( gene flow ). Despite the constant introduction of new variation through mutation and gene flow, most of
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#17328023256759048-458: A typical prokaryote and can be as much as a thousand times greater in volume. The main distinguishing feature of eukaryotes as compared to prokaryotes is compartmentalization : the presence of membrane-bound organelles (compartments) in which specific activities take place. Most important among these is a cell nucleus , an organelle that houses the cell's DNA . This nucleus gives the eukaryote its name, which means "true kernel (nucleus)". Some of
9222-454: Is directional selection , which is a shift in the average value of a trait over time—for example, organisms slowly getting taller. Secondly, disruptive selection is selection for extreme trait values and often results in two different values becoming most common, with selection against the average value. This would be when either short or tall organisms had an advantage, but not those of medium height. Finally, in stabilising selection there
9396-460: Is encoded in its DNA sequence. RNA is used for information transport (e.g., mRNA ) and enzymatic functions (e.g., ribosomal RNA). Transfer RNA (tRNA) molecules are used to add amino acids during protein translation . Prokaryotic genetic material is organized in a simple circular bacterial chromosome in the nucleoid region of the cytoplasm. Eukaryotic genetic material is divided into different, linear molecules called chromosomes inside
9570-442: Is more noticeable . Indeed, the evolution of microorganisms is particularly important to evolutionary research since their rapid reproduction allows the study of experimental evolution and the observation of evolution and adaptation in real time. Adaptation is the process that makes organisms better suited to their habitat. Also, the term adaptation may refer to a trait that is important for an organism's survival. For example,
9744-427: Is a byproduct of this process that may sometimes be adaptively beneficial. Gene flow is the exchange of genes between populations and between species. It can therefore be a source of variation that is new to a population or to a species. Gene flow can be caused by the movement of individuals between separate populations of organisms, as might be caused by the movement of mice between inland and coastal populations, or
9918-417: Is an inherited characteristic and an individual might inherit the "brown-eye trait" from one of their parents. Inherited traits are controlled by genes and the complete set of genes within an organism's genome (genetic material) is called its genotype . The complete set of observable traits that make up the structure and behaviour of an organism is called its phenotype . Some of these traits come from
10092-464: Is based on standing variation: when evolution depends on events of mutation that introduce new alleles, mutational and developmental biases in the introduction of variation (arrival biases) can impose biases on evolution without requiring neutral evolution or high mutation rates. Several studies report that the mutations implicated in adaptation reflect common mutation biases though others dispute this interpretation. Recombination allows alleles on
10266-407: Is called deep homology . During evolution, some structures may lose their original function and become vestigial structures. Such structures may have little or no function in a current species, yet have a clear function in ancestral species, or other closely related species. Examples include pseudogenes , the non-functional remains of eyes in blind cave-dwelling fish, wings in flightless birds,
10440-692: Is called genetic hitchhiking or genetic draft. Genetic draft caused by the fact that some neutral genes are genetically linked to others that are under selection can be partially captured by an appropriate effective population size. A special case of natural selection is sexual selection, which is selection for any trait that increases mating success by increasing the attractiveness of an organism to potential mates. Traits that evolved through sexual selection are particularly prominent among males of several animal species. Although sexually favoured, traits such as cumbersome antlers, mating calls, large body size and bright colours often attract predation, which compromises
10614-476: Is caused by a toxin released by the bacteria that grow from the spores. Clostridioides difficile infection , a common problem in healthcare settings, is caused by spore-forming bacteria. Bacteria exhibit an extremely wide variety of metabolic types. The distribution of metabolic traits within a group of bacteria has traditionally been used to define their taxonomy , but these traits often do not correspond with modern genetic classifications. Bacterial metabolism
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#173280232567510788-410: Is classified into nutritional groups on the basis of three major criteria: the source of energy , the electron donors used, and the source of carbon used for growth. Phototrophic bacteria derive energy from light using photosynthesis , while chemotrophic bacteria breaking down chemical compounds through oxidation , driving metabolism by transferring electrons from a given electron donor to
10962-534: Is dependent on bacterial secretion systems . These transfer proteins from the cytoplasm into the periplasm or into the environment around the cell. Many types of secretion systems are known and these structures are often essential for the virulence of pathogens, so are intensively studied. Some genera of Gram-positive bacteria, such as Bacillus , Clostridium , Sporohalobacter , Anaerobacter , and Heliobacterium , can form highly resistant, dormant structures called endospores . Endospores develop within
11136-697: Is detected. Diverse repair processes have evolved in organisms ranging from bacteria to humans. The widespread prevalence of these repair processes indicates the importance of maintaining cellular DNA in an undamaged state in order to avoid cell death or errors of replication due to damage that could lead to mutation . E. coli bacteria are a well-studied example of a cellular organism with diverse well-defined DNA repair processes. These include: nucleotide excision repair , DNA mismatch repair , non-homologous end joining of double-strand breaks, recombinational repair and light-dependent repair ( photoreactivation ). Between successive cell divisions, cells grow through
11310-532: Is determined by the bacterial cell wall and cytoskeleton and is important because it can influence the ability of bacteria to acquire nutrients, attach to surfaces, swim through liquids and escape predators . Multicellularity . Most bacterial species exist as single cells; others associate in characteristic patterns: Neisseria forms diploids (pairs), streptococci form chains, and staphylococci group together in "bunch of grapes" clusters. Bacteria can also group to form larger multicellular structures, such as
11484-435: Is different from that of eukaryotes and archaea. Some bacteria produce intracellular nutrient storage granules, such as glycogen , polyphosphate , sulfur or polyhydroxyalkanoates . Bacteria such as the photosynthetic cyanobacteria , produce internal gas vacuoles , which they use to regulate their buoyancy, allowing them to move up or down into water layers with different light intensities and nutrient levels. Around
11658-413: Is essential to the survival of many bacteria, and the antibiotic penicillin (produced by a fungus called Penicillium ) is able to kill bacteria by inhibiting a step in the synthesis of peptidoglycan. There are broadly speaking two different types of cell wall in bacteria, that classify bacteria into Gram-positive bacteria and Gram-negative bacteria . The names originate from the reaction of cells to
11832-597: Is from cyanobacteria -like organisms that lived between 3 and 3.5 billion years ago. Other early fossils of multicellular organisms include the contested Grypania spiralis and the fossils of the black shales of the Palaeoproterozoic Francevillian Group Fossil B Formation in Gabon . The evolution of multicellularity from unicellular ancestors has been replicated in the laboratory, in evolution experiments using predation as
12006-781: Is less well-studied but is involved in the maintenance of cell shape, polarity and cytokinesis. The subunit protein of microfilaments is a small, monomeric protein called actin . The subunit of microtubules is a dimeric molecule called tubulin . Intermediate filaments are heteropolymers whose subunits vary among the cell types in different tissues. Some of the subunit proteins of intermediate filaments include vimentin , desmin , lamin (lamins A, B and C), keratin (multiple acidic and basic keratins), and neurofilament proteins ( NF–L , NF–M ). Two different kinds of genetic material exist: deoxyribonucleic acid (DNA) and ribonucleic acid (RNA). Cells use DNA for their long-term information storage. The biological information contained in an organism
12180-527: Is made of about 20 proteins, with approximately another 30 proteins required for its regulation and assembly. The flagellum is a rotating structure driven by a reversible motor at the base that uses the electrochemical gradient across the membrane for power. Bacteria can use flagella in different ways to generate different kinds of movement. Many bacteria (such as E. coli ) have two distinct modes of movement: forward movement (swimming) and tumbling. The tumbling allows them to reorient and makes their movement
12354-466: Is made primarily of phospholipids . This membrane encloses the contents of the cell and acts as a barrier to hold nutrients, proteins and other essential components of the cytoplasm within the cell. Unlike eukaryotic cells , bacteria usually lack large membrane-bound structures in their cytoplasm such as a nucleus , mitochondria , chloroplasts and the other organelles present in eukaryotic cells. However, some bacteria have protein-bound organelles in
12528-594: Is measured by an organism's ability to survive and reproduce, which determines the size of its genetic contribution to the next generation. However, fitness is not the same as the total number of offspring: instead fitness is indicated by the proportion of subsequent generations that carry an organism's genes. For example, if an organism could survive well and reproduce rapidly, but its offspring were all too small and weak to survive, this organism would make little genetic contribution to future generations and would thus have low fitness. If an allele increases fitness more than
12702-494: Is motile in liquid or solid media. Several Listeria and Shigella species move inside host cells by usurping the cytoskeleton , which is normally used to move organelles inside the cell. By promoting actin polymerisation at one pole of their cells, they can form a kind of tail that pushes them through the host cell's cytoplasm. A few bacteria have chemical systems that generate light. This bioluminescence often occurs in bacteria that live in association with fish, and
12876-510: Is present in some bacteria outside the cell membrane and cell wall. The capsule may be polysaccharide as in pneumococci , meningococci or polypeptide as Bacillus anthracis or hyaluronic acid as in streptococci . Capsules are not marked by normal staining protocols and can be detected by India ink or methyl blue , which allows for higher contrast between the cells for observation. Flagella are organelles for cellular mobility. The bacterial flagellum stretches from cytoplasm through
13050-421: Is selection against extreme trait values on both ends, which causes a decrease in variance around the average value and less diversity. This would, for example, cause organisms to eventually have a similar height. Natural selection most generally makes nature the measure against which individuals and individual traits, are more or less likely to survive. "Nature" in this sense refers to an ecosystem , that is,
13224-429: Is that evolution has goals, long-term plans, or an innate tendency for "progress", as expressed in beliefs such as orthogenesis and evolutionism; realistically, however, evolution has no long-term goal and does not necessarily produce greater complexity. Although complex species have evolved, they occur as a side effect of the overall number of organisms increasing, and simple forms of life still remain more common in
13398-462: Is the stationary phase and is caused by depleted nutrients. The cells reduce their metabolic activity and consume non-essential cellular proteins. The stationary phase is a transition from rapid growth to a stress response state and there is increased expression of genes involved in DNA repair , antioxidant metabolism and nutrient transport . The final phase is the death phase where the bacteria run out of nutrients and die. Most bacteria have
13572-440: Is the logarithmic phase , also known as the exponential phase. The log phase is marked by rapid exponential growth . The rate at which cells grow during this phase is known as the growth rate ( k ), and the time it takes the cells to double is known as the generation time ( g ). During log phase, nutrients are metabolised at maximum speed until one of the nutrients is depleted and starts limiting growth. The third phase of growth
13746-447: Is the nearly neutral theory , according to which a mutation that would be effectively neutral in a small population is not necessarily neutral in a large population. Other theories propose that genetic drift is dwarfed by other stochastic forces in evolution, such as genetic hitchhiking, also known as genetic draft. Another concept is constructive neutral evolution (CNE), which explains that complex systems can emerge and spread into
13920-495: Is the more common means of reproduction among eukaryotes and multicellular organisms. The Red Queen hypothesis has been used to explain the significance of sexual reproduction as a means to enable continual evolution and adaptation in response to coevolution with other species in an ever-changing environment. Another hypothesis is that sexual reproduction is primarily an adaptation for promoting accurate recombinational repair of damage in germline DNA, and that increased diversity
14094-689: Is the plural of the Neo-Latin bacterium , which is the Latinisation of the Ancient Greek βακτήριον ( baktḗrion ), the diminutive of βακτηρία ( baktēría ), meaning "staff, cane", because the first ones to be discovered were rod-shaped . The ancestors of bacteria were unicellular microorganisms that were the first forms of life to appear on Earth, about 4 billion years ago. For about 3 billion years, most organisms were microscopic, and bacteria and archaea were
14268-439: Is the process by which traits that enhance survival and reproduction become more common in successive generations of a population. It embodies three principles: More offspring are produced than can possibly survive, and these conditions produce competition between organisms for survival and reproduction. Consequently, organisms with traits that give them an advantage over their competitors are more likely to pass on their traits to
14442-471: The Gram stain , a long-standing test for the classification of bacterial species. Gram-positive bacteria possess a thick cell wall containing many layers of peptidoglycan and teichoic acids . In contrast, Gram-negative bacteria have a relatively thin cell wall consisting of a few layers of peptidoglycan surrounded by a second lipid membrane containing lipopolysaccharides and lipoproteins . Most bacteria have
14616-600: The gut , though there are many on the skin. Most of the bacteria in and on the body are harmless or rendered so by the protective effects of the immune system , and many are beneficial , particularly the ones in the gut. However, several species of bacteria are pathogenic and cause infectious diseases , including cholera , syphilis , anthrax , leprosy , tuberculosis , tetanus and bubonic plague . The most common fatal bacterial diseases are respiratory infections . Antibiotics are used to treat bacterial infections and are also used in farming, making antibiotic resistance
14790-483: The heritable characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, resulting in certain characteristics becoming more or less common within a population over successive generations. The process of evolution has given rise to biodiversity at every level of biological organisation . The scientific theory of evolution by natural selection
14964-425: The neutral theory of molecular evolution most evolutionary changes are the result of the fixation of neutral mutations by genetic drift. In this model, most genetic changes in a population are thus the result of constant mutation pressure and genetic drift. This form of the neutral theory has been debated since it does not seem to fit some genetic variation seen in nature. A better-supported version of this model
15138-454: The product of a gene , or prevent the gene from functioning, or have no effect. About half of the mutations in the coding regions of protein-coding genes are deleterious — the other half are neutral. A small percentage of the total mutations in this region confer a fitness benefit. Some of the mutations in other parts of the genome are deleterious but the vast majority are neutral. A few are beneficial. Mutations can involve large sections of
15312-401: The selective pressure . The origin of cells has to do with the origin of life , which began the history of life on Earth. Small molecules needed for life may have been carried to Earth on meteorites, created at deep-sea vents , or synthesized by lightning in a reducing atmosphere . There is little experimental data defining what the first self-replicating forms were. RNA may have been
15486-442: The spirochaetes , are found between two membranes in the periplasmic space. They have a distinctive helical body that twists about as it moves. Two other types of bacterial motion are called twitching motility that relies on a structure called the type IV pilus , and gliding motility , that uses other mechanisms. In twitching motility, the rod-like pilus extends out from the cell, binds some substrate, and then retracts, pulling
15660-549: The vacuum and radiation of outer space , leading to the possibility that bacteria could be distributed throughout the Universe by space dust , meteoroids , asteroids , comets , planetoids , or directed panspermia . Endospore-forming bacteria can cause disease; for example, anthrax can be contracted by the inhalation of Bacillus anthracis endospores, and contamination of deep puncture wounds with Clostridium tetani endospores causes tetanus , which, like botulism ,
15834-565: 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 are parts of the cell that are adapted and/or specialized for carrying out one or more vital functions, analogous to the organs of the human body (such as the heart, lung, and kidney, with each organ performing a different function). Both eukaryotic and prokaryotic cells have organelles, but prokaryotic organelles are generally simpler and are not membrane-bound. There are several types of organelles in
16008-791: The Gram-negative cell wall, and only members of the Bacillota group and actinomycetota (previously known as the low G+C and high G+C Gram-positive bacteria, respectively) have the alternative Gram-positive arrangement. These differences in structure can produce differences in antibiotic susceptibility; for instance, vancomycin can kill only Gram-positive bacteria and is ineffective against Gram-negative pathogens , such as Haemophilus influenzae or Pseudomonas aeruginosa . Some bacteria have cell wall structures that are neither classically Gram-positive or Gram-negative. This includes clinically important bacteria such as mycobacteria which have
16182-467: The S phase of the cell cycle . In meiosis, the DNA is replicated only once, while the cell divides twice. DNA replication only occurs before meiosis I . DNA replication does not occur when the cells divide the second time, in meiosis II . Replication, like all cellular activities, requires specialized proteins for carrying out the job. Cells of all organisms contain enzyme systems that scan their DNA for damage and carry out repair processes when it
16356-519: The ability of organisms to generate genetic diversity and adapt by natural selection (increasing organisms' evolvability). Adaptation occurs through the gradual modification of existing structures. Consequently, structures with similar internal organisation may have different functions in related organisms. This is the result of a single ancestral structure being adapted to function in different ways. The bones within bat wings, for example, are very similar to those in mice feet and primate hands, due to
16530-402: The adaptation of horses' teeth to the grinding of grass. By using the term adaptation for the evolutionary process and adaptive trait for the product (the bodily part or function), the two senses of the word may be distinguished. Adaptations are produced by natural selection. The following definitions are due to Theodosius Dobzhansky: Adaptation may cause either the gain of a new feature, or
16704-502: The air, soil, water, acidic hot springs , radioactive waste , and the deep biosphere of Earth's crust . Bacteria play a vital role in many stages of the nutrient cycle by recycling nutrients and the fixation of nitrogen from the atmosphere . The nutrient cycle includes the decomposition of dead bodies ; bacteria are responsible for the putrefaction stage in this process. In the biological communities surrounding hydrothermal vents and cold seeps , extremophile bacteria provide
16878-507: The alleles are subject to sampling error . This drift halts when an allele eventually becomes fixed, either by disappearing from the population or by replacing the other alleles entirely. Genetic drift may therefore eliminate some alleles from a population due to chance alone. Even in the absence of selective forces, genetic drift can cause two separate populations that begin with the same genetic structure to drift apart into two divergent populations with different sets of alleles. According to
17052-418: The archaeal/eukaryotic lineage. The most recent common ancestor (MRCA) of bacteria and archaea was probably a hyperthermophile that lived about 2.5 billion–3.2 billion years ago. The earliest life on land may have been bacteria some 3.22 billion years ago. Bacteria were also involved in the second great evolutionary divergence, that of the archaea and eukaryotes. Here, eukaryotes resulted from
17226-452: The atmosphere and one cubic metre of air holds around one hundred million bacterial cells. The oceans and seas harbour around 3 x 10 bacteria which provide up to 50% of the oxygen humans breathe. Only around 2% of bacterial species have been fully studied. Size . Bacteria display a wide diversity of shapes and sizes. Bacterial cells are about one-tenth the size of eukaryotic cells and are typically 0.5–5.0 micrometres in length. However,
17400-488: The attachment of bacteria to specific receptors on human cells ( cell adhesion ). There are special types of pili involved in bacterial conjugation . Cell division involves a single cell (called a mother cell ) dividing into two daughter cells. This leads to growth in multicellular organisms (the growth of tissue ) and to procreation ( vegetative reproduction ) in unicellular organisms . Prokaryotic cells divide by binary fission , while eukaryotic cells usually undergo
17574-517: The bacteria have come into contact with in the past, which allows them to block virus replication through a form of RNA interference . Third, bacteria can transfer genetic material through direct cell contact via conjugation . In ordinary circumstances, transduction, conjugation, and transformation involve transfer of DNA between individual bacteria of the same species, but occasionally transfer may occur between individuals of different bacterial species, and this may have significant consequences, such as
17748-441: The bacteria perform separate tasks; for example, about one in ten cells migrate to the top of a fruiting body and differentiate into a specialised dormant state called a myxospore, which is more resistant to drying and other adverse environmental conditions. Biofilms . Bacteria often attach to surfaces and form dense aggregations called biofilms and larger formations known as microbial mats . These biofilms and mats can range from
17922-410: The bacterial chromosome, introducing foreign DNA in a process known as transduction . Many types of bacteriophage exist; some infect and lyse their host bacteria, while others insert into the bacterial chromosome. Bacteria resist phage infection through restriction modification systems that degrade foreign DNA and a system that uses CRISPR sequences to retain fragments of the genomes of phage that
18096-441: The biosphere. For example, the overwhelming majority of species are microscopic prokaryotes , which form about half the world's biomass despite their small size and constitute the vast majority of Earth's biodiversity. Simple organisms have therefore been the dominant form of life on Earth throughout its history and continue to be the main form of life up to the present day, with complex life only appearing more diverse because it
18270-432: The cell forward. Motile bacteria are attracted or repelled by certain stimuli in behaviours called taxes : these include chemotaxis , phototaxis , energy taxis , and magnetotaxis . In one peculiar group, the myxobacteria, individual bacteria move together to form waves of cells that then differentiate to form fruiting bodies containing spores. The myxobacteria move only when on solid surfaces, unlike E. coli , which
18444-416: The cell in processes of growth and mobility. The eukaryotic cytoskeleton is composed of microtubules , intermediate filaments and microfilaments . In the cytoskeleton of a neuron the intermediate filaments are known as neurofilaments . There are a great number of proteins associated with them, each controlling a cell's structure by directing, bundling, and aligning filaments. The prokaryotic cytoskeleton
18618-444: The cell membrane(s) and extrudes through the cell wall. They are long and thick thread-like appendages, protein in nature. A different type of flagellum is found in archaea and a different type is found in eukaryotes. A fimbria (plural fimbriae also known as a pilus , plural pili) is a short, thin, hair-like filament found on the surface of bacteria. Fimbriae are formed of a protein called pilin ( antigenic ) and are responsible for
18792-449: The cell, glucose is broken down to make adenosine triphosphate ( ATP ), a molecule that possesses readily available energy, through two different pathways. In plant cells, chloroplasts create sugars by photosynthesis , using the energy of light to join molecules of water and carbon dioxide . Cells are capable of synthesizing new proteins, which are essential for the modulation and maintenance of cellular activities. This process involves
18966-436: 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
19140-488: The change over time in this genetic variation. The frequency of one particular allele will become more or less prevalent relative to other forms of that gene. Variation disappears when a new allele reaches the point of fixation —when it either disappears from the population or replaces the ancestral allele entirely. Mutations are changes in the DNA sequence of a cell's genome and are the ultimate source of genetic variation in all organisms. When mutations occur, they may alter
19314-575: The class Schizomycetes ("fission fungi"), bacteria are now classified as prokaryotes . Unlike cells of animals and other eukaryotes , bacterial cells do not contain a nucleus and rarely harbour membrane -bound organelles . Although the term bacteria traditionally included all prokaryotes, the scientific classification changed after the discovery in the 1990s that prokaryotes consist of two very different groups of organisms that evolved from an ancient common ancestor . These evolutionary domains are called Bacteria and Archaea . The word bacteria
19488-623: The cytoplasm of the cell; generally, a single endospore develops in each cell. Each endospore contains a core of DNA and ribosomes surrounded by a cortex layer and protected by a multilayer rigid coat composed of peptidoglycan and a variety of proteins. Endospores show no detectable metabolism and can survive extreme physical and chemical stresses, such as high levels of UV light , gamma radiation , detergents , disinfectants , heat, freezing, pressure, and desiccation . In this dormant state, these organisms may remain viable for millions of years. Endospores even allow bacteria to survive exposure to
19662-413: The cytoplasm which compartmentalise aspects of bacterial metabolism, such as the carboxysome . Additionally, bacteria have a multi-component cytoskeleton to control the localisation of proteins and nucleic acids within the cell, and to manage the process of cell division . Many important biochemical reactions, such as energy generation, occur due to concentration gradients across membranes, creating
19836-911: The cytoskeleton. In August 2020, scientists described one way cells—in particular cells of a slime mold and mouse pancreatic cancer-derived cells—are able to navigate efficiently through a body and identify the best routes through complex mazes: generating gradients after breaking down diffused chemoattractants which enable them to sense upcoming maze junctions before reaching them, including around corners. Multicellular organisms are organisms that consist of more than one cell, in contrast to single-celled organisms . In complex multicellular organisms, cells specialize into different cell types that are adapted to particular functions. In mammals, major cell types include skin cells , muscle cells , neurons , blood cells , fibroblasts , stem cells , and others. Cell types differ both in appearance and function, yet are genetically identical. Cells are able to be of
20010-403: The descent of all these structures from a common mammalian ancestor. However, since all living organisms are related to some extent, even organs that appear to have little or no structural similarity, such as arthropod , squid and vertebrate eyes, or the limbs and wings of arthropods and vertebrates, can depend on a common set of homologous genes that control their assembly and function; this
20184-400: The development of thinking about the evolution of genome composition, including isochores. Different insertion vs. deletion biases in different taxa can lead to the evolution of different genome sizes. The hypothesis of Lynch regarding genome size relies on mutational biases toward increase or decrease in genome size. However, mutational hypotheses for the evolution of composition suffered
20358-490: The direct control of genes include the inheritance of cultural traits and symbiogenesis . From a neo-Darwinian perspective, evolution occurs when there are changes in the frequencies of alleles within a population of interbreeding organisms, for example, the allele for black colour in a population of moths becoming more common. Mechanisms that can lead to changes in allele frequencies include natural selection, genetic drift, and mutation bias. Evolution by natural selection
20532-408: The dominant forms of life. Although bacterial fossils exist, such as stromatolites , their lack of distinctive morphology prevents them from being used to examine the history of bacterial evolution, or to date the time of origin of a particular bacterial species. However, gene sequences can be used to reconstruct the bacterial phylogeny , and these studies indicate that bacteria diverged first from
20706-493: The earliest self-replicating molecule , as it can both store genetic information and catalyze chemical reactions. Cells emerged around 4 billion years ago. The first cells were most likely heterotrophs . The early cell membranes were probably simpler and more permeable than modern ones, with only a single fatty acid chain per lipid. Lipids spontaneously form bilayered vesicles in water, and could have preceded RNA. Eukaryotic cells were created some 2.2 billion years ago in
20880-524: The elongated filaments of Actinomycetota species, the aggregates of Myxobacteria species, and the complex hyphae of Streptomyces species. These multicellular structures are often only seen in certain conditions. For example, when starved of amino acids, myxobacteria detect surrounding cells in a process known as quorum sensing , migrate towards each other, and aggregate to form fruiting bodies up to 500 micrometres long and containing approximately 100,000 bacterial cells. In these fruiting bodies,
21054-499: The energy released by the transfer of ions down an electrochemical gradient across the cell membrane. Fimbriae (sometimes called " attachment pili ") are fine filaments of protein, usually 2–10 nanometres in diameter and up to several micrometres in length. They are distributed over the surface of the cell, and resemble fine hairs when seen under the electron microscope . Fimbriae are believed to be involved in attachment to solid surfaces or to other cells, and are essential for
21228-600: The entering of ancient bacteria into endosymbiotic associations with the ancestors of eukaryotic cells, which were themselves possibly related to the Archaea. This involved the engulfment by proto-eukaryotic cells of alphaproteobacterial symbionts to form either mitochondria or hydrogenosomes , which are still found in all known Eukarya (sometimes in highly reduced form , e.g. in ancient "amitochondrial" protozoa). Later, some eukaryotes that already contained mitochondria also engulfed cyanobacteria -like organisms, leading to
21402-638: The evolution of different growth strategies (see r/K selection theory ). Some organisms can grow extremely rapidly when nutrients become available, such as the formation of algal and cyanobacterial blooms that often occur in lakes during the summer. Other organisms have adaptations to harsh environments, such as the production of multiple antibiotics by Streptomyces that inhibit the growth of competing microorganisms. In nature, many organisms live in communities (e.g., biofilms ) that may allow for increased supply of nutrients and protection from environmental stresses. These relationships can be essential for growth of
21576-489: The evolutionary history of life on Earth. Morphological and biochemical traits tend to be more similar among species that share a more recent common ancestor , which historically was used to reconstruct phylogenetic trees , although direct comparison of genetic sequences is a more common method today. Evolutionary biologists have continued to study various aspects of evolution by forming and testing hypotheses as well as constructing theories based on evidence from
21750-633: The external environment by the cell membrane. In order to assemble these structures, their components must be carried across the cell membrane by export processes. Many types of prokaryotic and eukaryotic cells have a cell wall . The cell wall acts to protect the cell mechanically and chemically from its environment, and is an additional layer of protection to the cell membrane. Different types of cell have cell walls made up of different materials; plant cell walls are primarily made up of cellulose , fungi cell walls are made up of chitin and bacteria cell walls are made up of peptidoglycan . A gelatinous capsule
21924-414: 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 also other fields including agriculture, medicine, and computer science . Evolution in organisms occurs through changes in heritable characteristics—the inherited characteristics of an organism. In humans, for example, eye colour
22098-427: The fitness of an allele is not a fixed characteristic; if the environment changes, previously neutral or harmful traits may become beneficial and previously beneficial traits become harmful. However, even if the direction of selection does reverse in this way, traits that were lost in the past may not re-evolve in an identical form. However, a re-activation of dormant genes, as long as they have not been eliminated from
22272-563: The formation of chloroplasts in algae and plants. This is known as primary endosymbiosis . Bacteria are ubiquitous, living in every possible habitat on the planet including soil, underwater, deep in Earth's crust and even such extreme environments as acidic hot springs and radioactive waste. There are thought to be approximately 2×10 bacteria on Earth, forming a biomass that is only exceeded by plants. They are abundant in lakes and oceans, in arctic ice, and geothermal springs where they provide
22446-470: 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
22620-523: 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 can be broken down into simpler sugar molecules called monosaccharides such as glucose . Once inside
22794-472: The genome and were only suppressed perhaps for hundreds of generations, can lead to the re-occurrence of traits thought to be lost like hindlegs in dolphins, teeth in chickens, wings in wingless stick insects, tails and additional nipples in humans etc. "Throwbacks" such as these are known as atavisms . Natural selection within a population for a trait that can vary across a range of values, such as height, can be categorised into three different types. The first
22968-479: The genome of a species is very similar among all individuals of that species. However, discoveries in the field of evolutionary developmental biology have demonstrated that even relatively small differences in genotype can lead to dramatic differences in phenotype both within and between species. An individual organism's phenotype results from both its genotype and the influence of the environment it has lived in. The modern evolutionary synthesis defines evolution as
23142-650: The growth in cell population. Cell (biology) The cell is the basic structural and functional unit of all forms of life . Every cell consists of cytoplasm enclosed within a membrane ; many cells contain organelles , each with a specific function. The term comes from the Latin word cellula meaning 'small room'. Most cells are only visible under a microscope . Cells emerged on Earth about 4 billion years ago. All cells are capable of replication , protein synthesis , and motility . Cells are broadly categorized into two types: eukaryotic cells , which possess
23316-640: The idea of developmental bias . Haldane and Fisher argued that, because mutation is a weak pressure easily overcome by selection, tendencies of mutation would be ineffectual except under conditions of neutral evolution or extraordinarily high mutation rates. This opposing-pressures argument was long used to dismiss the possibility of internal tendencies in evolution, until the molecular era prompted renewed interest in neutral evolution. Noboru Sueoka and Ernst Freese proposed that systematic biases in mutation might be responsible for systematic differences in genomic GC composition between species. The identification of
23490-399: The information necessary to build various proteins such as enzymes , the cell's primary machinery. There are also other kinds of biomolecules in cells. This article lists these primary cellular components , then briefly describes their function. The cell membrane , or plasma membrane, is a selectively permeable biological membrane that surrounds the cytoplasm of a cell. In animals,
23664-450: The interaction of its genotype with the environment while others are neutral. Some observable characteristics are not inherited. For example, suntanned skin comes from the interaction between a person's genotype and sunlight; thus, suntans are not passed on to people's children. The phenotype is the ability of the skin to tan when exposed to sunlight. However, some people tan more easily than others, due to differences in genotypic variation;
23838-623: The largest viruses . Some bacteria may be even smaller, but these ultramicrobacteria are not well-studied. Shape . Most bacterial species are either spherical, called cocci ( singular coccus , from Greek kókkos , grain, seed), or rod-shaped, called bacilli ( sing . bacillus, from Latin baculus , stick). Some bacteria, called vibrio , are shaped like slightly curved rods or comma-shaped; others can be spiral-shaped, called spirilla , or tightly coiled, called spirochaetes . A small number of other unusual shapes have been described, such as star-shaped bacteria. This wide variety of shapes
24012-512: The level of the individual organism are genes called transposons , which can replicate and spread throughout a genome. Selection at a level above the individual, such as group selection , may allow the evolution of cooperation. Genetic drift is the random fluctuation of allele frequencies within a population from one generation to the next. When selective forces are absent or relatively weak, allele frequencies are equally likely to drift upward or downward in each successive generation because
24186-712: The light probably serves to attract fish or other large animals. Bacteria often function as multicellular aggregates known as biofilms , exchanging a variety of molecular signals for intercell communication and engaging in coordinated multicellular behaviour. The communal benefits of multicellular cooperation include a cellular division of labour , accessing resources that cannot effectively be used by single cells, collectively defending against antagonists, and optimising population survival by differentiating into distinct cell types. For example, bacteria in biofilms can have more than five hundred times increased resistance to antibacterial agents than individual "planktonic" bacteria of
24360-452: The longer term, evolution produces new species through splitting ancestral populations of organisms into new groups that cannot or will not interbreed. These outcomes of evolution are distinguished based on time scale as macroevolution versus microevolution. Macroevolution refers to evolution that occurs at or above the level of species, in particular speciation and extinction, whereas microevolution refers to smaller evolutionary changes within
24534-405: The loss of an ancestral feature. An example that shows both types of change is bacterial adaptation to antibiotic selection, with genetic changes causing antibiotic resistance by both modifying the target of the drug, or increasing the activity of transporters that pump the drug out of the cell. Other striking examples are the bacteria Escherichia coli evolving the ability to use citric acid as
24708-458: The mechanics in developmental plasticity and canalisation . Heritability may also occur at even larger scales. For example, ecological inheritance through the process of niche construction is defined by the regular and repeated activities of organisms in their environment. This generates a legacy of effects that modify and feed back into the selection regime of subsequent generations. Other examples of heritability in evolution that are not under
24882-432: The microorganisms that cause infection. Cell motility involves many receptors, crosslinking, bundling, binding, adhesion, motor and other proteins. 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
25056-408: The movement of pollen between heavy-metal-tolerant and heavy-metal-sensitive populations of grasses. Gene transfer between species includes the formation of hybrid organisms and horizontal gene transfer . Horizontal gene transfer is the transfer of genetic material from one organism to another organism that is not its offspring; this is most common among bacteria. In medicine, this contributes to
25230-415: The newly formed daughter cells. Examples include fruiting body formation by myxobacteria and aerial hyphae formation by Streptomyces species, or budding. Budding involves a cell forming a protrusion that breaks away and produces a daughter cell. In the laboratory, bacteria are usually grown using solid or liquid media. Solid growth media , such as agar plates , are used to isolate pure cultures of
25404-427: The next generation than those with traits that do not confer an advantage. This teleonomy is the quality whereby the process of natural selection creates and preserves traits that are seemingly fitted for the functional roles they perform. Consequences of selection include nonrandom mating and genetic hitchhiking . The central concept of natural selection is the evolutionary fitness of an organism. Fitness
25578-514: The nutrients needed to sustain life by converting dissolved compounds, such as hydrogen sulphide and methane , to energy. Bacteria also live in mutualistic , commensal and parasitic relationships with plants and animals. Most bacteria have not been characterised and there are many species that cannot be grown in the laboratory. The study of bacteria is known as bacteriology , a branch of microbiology . Like all animals, humans carry vast numbers (approximately 10 to 10) of bacteria. Most are in
25752-470: The nutrients needed to sustain life by converting dissolved compounds, such as hydrogen sulphide and methane , to energy. They live on and in plants and animals. Most do not cause diseases, are beneficial to their environments, and are essential for life. The soil is a rich source of bacteria and a few grams contain around a thousand million of them. They are all essential to soil ecology, breaking down toxic waste and recycling nutrients. They are even found in
25926-499: The offspring of sexual organisms contain random mixtures of their parents' chromosomes that are produced through independent assortment. In a related process called homologous recombination , sexual organisms exchange DNA between two matching chromosomes. Recombination and reassortment do not alter allele frequencies, but instead change which alleles are associated with each other, producing offspring with new combinations of alleles. Sex usually increases genetic variation and may increase
26100-456: The other alleles of that gene, then with each generation this allele has a higher probability of becoming common within the population. These traits are said to be "selected for ." Examples of traits that can increase fitness are enhanced survival and increased fecundity . Conversely, the lower fitness caused by having a less beneficial or deleterious allele results in this allele likely becoming rarer—they are "selected against ." Importantly,
26274-455: The other differences are: Many groups of eukaryotes are single-celled. Among the many-celled groups are animals and plants. The number of cells in these groups vary with species; it has been estimated that the human body contains around 37 trillion (3.72×10 ) cells, and more recent studies put this number at around 30 trillion (~36 trillion cells in the male, ~28 trillion in the female). All cells, whether prokaryotic or eukaryotic , have
26448-473: The outside of the cell membrane is the cell wall . Bacterial cell walls are made of peptidoglycan (also called murein), which is made from polysaccharide chains cross-linked by peptides containing D- amino acids . Bacterial cell walls are different from the cell walls of plants and fungi , which are made of cellulose and chitin , respectively. The cell wall of bacteria is also distinct from that of achaea, which do not contain peptidoglycan. The cell wall
26622-578: The overall process, like a step in an assembly line. One example of mutation is wild boar piglets. They are camouflage coloured and show a characteristic pattern of dark and light longitudinal stripes. However, mutations in the melanocortin 1 receptor ( MC1R ) disrupt the pattern. The majority of pig breeds carry MC1R mutations disrupting wild-type colour and different mutations causing dominant black colouring. In asexual organisms, genes are inherited together, or linked , as they cannot mix with genes of other organisms during reproduction. In contrast,
26796-409: The plasma membrane is the outer boundary of the cell, while in plants and prokaryotes it is usually covered by a cell wall . This membrane serves to separate and protect a cell from its surrounding environment and is made mostly from a double layer of phospholipids , which are amphiphilic (partly hydrophobic and partly hydrophilic ). Hence, the layer is called a phospholipid bilayer , or sometimes
26970-527: The presence of hip bones in whales and snakes, and sexual traits in organisms that reproduce via asexual reproduction. Examples of vestigial structures in humans include wisdom teeth , the coccyx , the vermiform appendix , and other behavioural vestiges such as goose bumps and primitive reflexes . However, many traits that appear to be simple adaptations are in fact exaptations : structures originally adapted for one function, but which coincidentally became somewhat useful for some other function in
27144-469: The process. One example is the African lizard Holaspis guentheri , which developed an extremely flat head for hiding in crevices, as can be seen by looking at its near relatives. However, in this species, the head has become so flattened that it assists in gliding from tree to tree—an exaptation. Within cells, molecular machines such as the bacterial flagella and protein sorting machinery evolved by
27318-610: The production of the highly toxic forms of mercury ( methyl- and dimethylmercury ) in the environment. Nonrespiratory anaerobes use fermentation to generate energy and reducing power, secreting metabolic by-products (such as ethanol in brewing) as waste. Facultative anaerobes can switch between fermentation and different terminal electron acceptors depending on the environmental conditions in which they find themselves. Unlike in multicellular organisms, increases in cell size ( cell growth ) and reproduction by cell division are tightly linked in unicellular organisms. Bacteria grow to
27492-494: The rate of evolution. The two-fold cost of sex was first described by John Maynard Smith . The first cost is that in sexually dimorphic species only one of the two sexes can bear young. This cost does not apply to hermaphroditic species, like most plants and many invertebrates . The second cost is that any individual who reproduces sexually can only pass on 50% of its genes to any individual offspring, with even less passed on as each new generation passes. Yet sexual reproduction
27666-408: The relative importance of selection and neutral processes, including drift. The comparative importance of adaptive and non-adaptive forces in driving evolutionary change is an area of current research . Mutation bias is usually conceived as a difference in expected rates for two different kinds of mutation, e.g., transition-transversion bias, GC-AT bias, deletion-insertion bias. This is related to
27840-558: The replication of DNA or from exposure to mutagens . Mutation rates vary widely among different species of bacteria and even among different clones of a single species of bacteria. Genetic changes in bacterial genomes emerge from either random mutation during replication or "stress-directed mutation", where genes involved in a particular growth-limiting process have an increased mutation rate. Some bacteria transfer genetic material between cells. This can occur in three main ways. First, bacteria can take up exogenous DNA from their environment in
28014-443: The same genotype but of different cell type due to the differential expression of the genes they contain. Most distinct cell types arise from a single totipotent cell, called a zygote , that differentiates into hundreds of different cell types during the course of development . Differentiation of cells is driven by different environmental cues (such as cell–cell interaction) and intrinsic differences (such as those caused by
28188-527: The same species. One type of intercellular communication by a molecular signal is called quorum sensing , which serves the purpose of determining whether the local population density is sufficient to support investment in processes that are only successful if large numbers of similar organisms behave similarly, such as excreting digestive enzymes or emitting light. Quorum sensing enables bacteria to coordinate gene expression and to produce, release, and detect autoinducers or pheromones that accumulate with
28362-417: The same strand of DNA to become separated. However, the rate of recombination is low (approximately two events per chromosome per generation). As a result, genes close together on a chromosome may not always be shuffled away from each other and genes that are close together tend to be inherited together, a phenomenon known as linkage . This tendency is measured by finding how often two alleles occur together on
28536-552: The sequence of nucleotides in the DNA. These phenomena are classed as epigenetic inheritance systems. DNA methylation marking chromatin , self-sustaining metabolic loops, gene silencing by RNA interference and the three-dimensional conformation of proteins (such as prions ) are areas where epigenetic inheritance systems have been discovered at the organismic level. Developmental biologists suggest that complex interactions in genetic networks and communication among cells can lead to heritable variations that may underlay some of
28710-406: The spread of antibiotic resistance , as when one bacteria acquires resistance genes it can rapidly transfer them to other species. Horizontal transfer of genes from bacteria to eukaryotes such as the yeast Saccharomyces cerevisiae and the adzuki bean weevil Callosobruchus chinensis has occurred. An example of larger-scale transfers are the eukaryotic bdelloid rotifers , which have received
28884-410: The surfaces of plants, the majority of bacteria are bound to surfaces in biofilms. Biofilms are also important in medicine, as these structures are often present during chronic bacterial infections or in infections of implanted medical devices , and bacteria protected within biofilms are much harder to kill than individual isolated bacteria. The bacterial cell is surrounded by a cell membrane , which
29058-640: The survival of individual males. This survival disadvantage is balanced by higher reproductive success in males that show these hard-to-fake , sexually selected traits. Evolution influences every aspect of the form and behaviour of organisms. Most prominent are the specific behavioural and physical adaptations that are the outcome of natural selection. These adaptations increase fitness by aiding activities such as finding food, avoiding predators or attracting mates. Organisms can also respond to selection by cooperating with each other, usually by aiding their relatives or engaging in mutually beneficial symbiosis . In
29232-642: The system...." Each population within an ecosystem occupies a distinct niche , or position, with distinct relationships to other parts of the system. These relationships involve the life history of the organism, its position in the food chain and its geographic range. This broad understanding of nature enables scientists to delineate specific forces which, together, comprise natural selection. Natural selection can act at different levels of organisation , such as genes, cells, individual organisms, groups of organisms and species. Selection can act at multiple levels simultaneously. An example of selection occurring below
29406-455: The terminal electron acceptor, while anaerobic organisms use other compounds such as nitrate , sulfate , or carbon dioxide. Many bacteria, called heterotrophs , derive their carbon from other organic carbon . Others, such as cyanobacteria and some purple bacteria , are autotrophic , meaning they obtain cellular carbon by fixing carbon dioxide . In unusual circumstances, the gas methane can be used by methanotrophic bacteria as both
29580-443: The transfer of antibiotic resistance. In such cases, gene acquisition from other bacteria or the environment is called horizontal gene transfer and may be common under natural conditions. Many bacteria are motile (able to move themselves) and do so using a variety of mechanisms. The best studied of these are flagella , long filaments that are turned by a motor at the base to generate propeller-like movement. The bacterial flagellum
29754-748: The uneven distribution of molecules during division ). Multicellularity has evolved independently at least 25 times, including in some prokaryotes, like cyanobacteria , myxobacteria , actinomycetes , or Methanosarcina . However, complex multicellular organisms evolved only in six eukaryotic groups: animals, fungi, brown algae, red algae, green algae, and plants. It evolved repeatedly for plants ( Chloroplastida ), once or twice for animals , once for brown algae , and perhaps several times for fungi , slime molds , and red algae . Multicellularity may have evolved from colonies of interdependent organisms, from cellularization , or from organisms in symbiotic relationships . The first evidence of multicellularity
29928-491: The virulence of some bacterial pathogens. Pili ( sing . pilus) are cellular appendages, slightly larger than fimbriae, that can transfer genetic material between bacterial cells in a process called conjugation where they are called conjugation pili or sex pili (see bacterial genetics, below). They can also generate movement where they are called type IV pili . Glycocalyx is produced by many bacteria to surround their cells, and varies in structural complexity: ranging from
30102-434: The way, though the sequence of the steps involved has been disputed, and may not have started with symbiogenesis. It featured at least one centriole and cilium , sex ( meiosis and syngamy ), peroxisomes , and a dormant cyst with a cell wall of chitin and/or cellulose . In turn, the last eukaryotic common ancestor gave rise to the eukaryotes' crown group , containing the ancestors of animals , fungi , plants , and
30276-895: Was conceived independently by two British naturalists, Charles Darwin and Alfred Russel Wallace , in the mid-19th century as an explanation for why organisms are adapted to their physical and biological environments. The theory was first set out in detail in Darwin's book On the Origin of Species . Evolution by natural selection is established by observable facts about living organisms: (1) more offspring are often produced than can possibly survive; (2) traits vary among individuals with respect to their morphology , physiology , and behaviour; (3) different traits confer different rates of survival and reproduction (differential fitness ); and (4) traits can be passed from generation to generation ( heritability of fitness). In successive generations, members of
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