Chemotaxis - wikipedia Chemotaxis This article has multiple issues. Please help improve it or discuss these issues on the talk page. ( Learn how and when to remove these template messages ) This article needs additional citations for verification. Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed. ( September 2015 ) This article relies too much on references to primary sources. Please improve this by adding secondary or tertiary sources. ( March 2017 ) This article needs attention from an expert in Physiology. The specific problem is : the article was begun as an academic enterprise, with great expertise brought to bear, but was unfortunately developed without WP : VERIFY being followed. With the departure of the original academic team, the article is evolving with editorial additions where it seems sourcing is viewed as optional. The result is that : ( i ) original material still fails the test of being secondary source - based, ( ii ) completely independent perspectives on the research area covered in the original article, and since, is lacking, ( iii ) article updates are likewise not not based on secondary sources, and so there seems to be overall regression of the article over time. WikiProject Physiology may be able to help recruit an expert. ( March 2017 ) This article includes inline citations, but they are not properly formatted. Please improve this article by correcting them. ( March 2017 ) Capillary tube assay for chemotaxis. Motile prokaryotes sense chemicals in their environment and change their motility accordingly. Absent chemicals, movement is completely random. When an attractant or repellent is present, runs become longer and tumbles become less frequent. The result is net movement towards or away from the chemical ( i.e., up up or down the chemical gradient ). The net movement can be seen in the beaker, where the bacteria accumulate around the origin of the attractant, and away from the origin of the repellent. Chemotaxis ( from chemo - + taxis ) is the movement of an organism in response to a chemical stimulus. Somatic cells, bacteria, and other single - cell or multicellular organisms direct their movements according to certain chemicals in their environment. This is important for bacteria to find food ( e.g., glucose ) by swimming toward the highest concentration of food molecules, or to flee from poisons ( e.g., phenol ). In multicellular organisms, chemotaxis is critical to early development ( e.g., movement of sperm towards the egg during fertilization ) and subsequent phases of development ( e.g., migration of neurons or lymphocytes ) as well as in normal function. In addition, it has been recognized that mechanisms that allow chemotaxis in animals can be subverted during cancer metastasis. Positive chemotaxis occurs if the movement is toward a higher concentration of the chemical in question ; negative chemotaxis if the movement is in the opposite direction. Chemically prompted kinesis ( randomly directed or nondirectional ) can be called chemokinesis. Contents 1 History of chemotaxis research 2 Chemoattractants and chemorepellents 3 Bacterial chemotaxis -- general characteristics 3.1 Behavior 3.2 Signal transduction 3.2. 1 Flagellum regulation 3.2. 2 Receptor regulation 4 Eukaryotic chemotaxis 4.1 Motility 4.1. 1 Chemotaxis related migratory responses 4.2 Receptors 4.2. 1 Chemotactic selection 4.3 Chemotactic ligands 4.3. 1 Chemotactic range fitting 5 Clinical significance 6 Mathematical models 7 Measurement of chemotaxis 8 Artificial chemotactic systems 9 See also 10 References 11 Further reading 12 External links History of chemotaxis research This section needs additional citations for verification. Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed. ( March 2017 ) Although migration of cells was detected from the early days of the development of microscopy by Leeuwenhoek, a Caltech lecture regarding chemotaxis propounds that ' erudite description of chemotaxis was only first made by T.W. Engelmann ( 1881 ) and W.F. Pfeffer ( 1884 ) in bacteria, and H.S. Jennings ( 1906 ) in ciliates '. The Nobel Prize laureate I. Metchnikoff also contributed to the study of the field during 1882 to 1886, with investigations of the process as an initial step of phagocytosis. The significance of chemotaxis in biology and clinical pathology was widely accepted in the 1930s, and the most fundamental definitions underlying the phenomenon were drafted by this time. The most important aspects in quality control of chemotaxis assays were described by H. Harris in the 1950s. In the 1960s and 1970s, the revolution of modern cell biology and biochemistry provided a series of novel techniques that became available to investigate the migratory responder cells and subcellular fractions responsible for chemotactic activity. The availability of this technology led to the discovery of C5a, a major chemotactic factor involved in acute inflammation. The pioneering works of J. Adler represented a significant turning point in understanding the whole process of intracellular signal transduction of bacteria. Chemoattractants and chemorepellents This section has multiple issues. Please help improve it or discuss these issues on the talk page. ( Learn how and when to remove these template messages ) This section does not cite any sources. Please help improve this section by adding citations to reliable sources. Unsourced material may be challenged and removed. ( March 2017 ) This section needs expansion with : with an updated perspective, sufficiently complete that it clearly differentiates between procaryotic and eukaryotic processes in a way that young readers can understand, and that captures the different ways that the general process is used in each kingdom. You can help by adding to it. ( March 2017 ) Chemoattractants and chemorepellents are inorganic or organic substances possessing chemotaxis - inducer effect in motile cells. Effects of chemoattractants are elicited via described or hypothetic chemotaxis receptors, the chemoattractant moiety of a ligand is target cell specific and concentration dependent. Most frequently investigated chemoattractants are formyl peptides and chemokines. Responses to chemorepellents result in axial swimming and they are considered a basic motile phenomena in bacteria. The most frequently investigated chemorepellents are inorganic salts, amino acids, and some chemokines. Bacterial chemotaxis -- general characteristics This section has multiple issues. Please help improve it or discuss these issues on the talk page. ( Learn how and when to remove these template messages ) This section needs expansion with : with updated perspective that also makes clear what of the general model developed from studies in E. coli and S. enterica broadly applies, cf. Rao et al ( 2004 ), in Further reading, and articles since. You can help by adding to it. ( March 2017 ) This section needs additional citations for verification. Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed. ( March 2017 ) Some bacteria, such as E. coli, have several flagella per cell ( 4 -- 10 typically ). These can rotate in two ways : Counter-clockwise rotation aligns the flagella into a single rotating bundle, causing the bacterium to swim in a straight line ; and Clockwise rotation breaks the flagella bundle apart such that each flagellum points in a different direction, causing the bacterium to tumble in place. The directions of rotation are given for an observer outside the cell looking down the flagella toward the cell. Behavior The overall movement of a bacterium is the result of alternating tumble and swim phases. If one watches a bacterium swimming in a uniform environment, its movement will look like a random walk with relatively straight swims interrupted by random tumbles that reorient the bacterium. Bacteria such as E. coli are unable to choose the direction in which they swim, and are unable to swim in a straight line for more than a few seconds due to rotational diffusion ; in other words, bacteria `` forget '' the direction in which they are going. By repeatedly evaluating their course, and adjusting if they are moving in the wrong direction, bacteria can direct their motion to find favorable locations with high concentrations of attractants ( usually food ) and avoid repellents ( usually poisons ). In the presence of a chemical gradient bacteria will chemotax, or direct their overall motion based on the gradient. If the bacterium senses that it is moving in the correct direction ( toward attractant / away from repellent ), it will keep swimming in a straight line for a longer time before tumbling ; however, if it is moving in the wrong direction, it will tumble sooner and try a new direction at random. In other words, bacteria like E. coli use temporal sensing to decide whether their situation is improving or not, and in this way, find the location with the highest concentration of attractant ( usually the source ) quite well. Even under very high concentrations, it can still distinguish very small differences in concentration, and fleeing from a repellent works with the same efficiency. This biased random walk is a result of simply choosing between two methods of random movement ; namely tumbling and straight swimming. In fact, chemotactic responses such as forgetting direction and choosing movements resemble the decision - making abilities of higher life - forms with brains that process sensory data. The helical nature of the individual flagellar filament is critical for this movement to occur, and the protein that makes up the flagellar filament, flagellin, is quite similar among all flagellated bacteria. Vertebrates seem to have taken advantage of this fact by possessing an immune receptor ( TLR5 ) designed to recognize this conserved protein. As in many instances in biology, there are bacteria that do not follow this rule. Many bacteria, such as Vibrio, are monoflagellated and have a single flagellum at one pole of the cell. Their method of chemotaxis is different. Others possess a single flagellum that is kept inside the cell wall. These bacteria move by spinning the whole cell, which is shaped like a corkscrew. Signal transduction Chemical gradients are sensed through multiple transmembrane receptors, called methyl - accepting chemotaxis proteins ( MCPs ), which vary in the molecules that they detect. These receptors may bind attractants or repellents directly or indirectly through interaction with proteins of periplasmatic space. The signals from these receptors are transmitted across the plasma membrane into the cytosol, where Che proteins are activated. The Che proteins alter the tumbling frequency, and alter the receptors. Flagellum regulation The proteins CheW and CheA bind to the receptor. The absence of receptor activation results in autophosphorylation in the histidine kinase, CheA, at a single highly conserved histidine residue. CheA, in turn, transfers phosphoryl groups to conserved aspartate residues in the response regulators CheB and CheY ; CheA is a histidine kinase and it does not actively transfer the phosphoryl group, rather, the response regulator CheB takes the phosphoryl group from CheA. This mechanism of signal transduction is called a two - component system, and it is a common form of signal transduction in bacteria. CheY induces tumbling by interacting with the flagellar switch protein FliM, inducing a change from counter-clockwise to clockwise rotation of the flagellum. Change in the rotation state of a single flagellum can disrupt the entire flagella bundle and cause a tumble. Receptor regulation CheB, when activated by CheA, acts as a methylesterase, removing methyl groups from glutamate residues on the cytosolic side of the receptor ; it works antagonistically with CheR, a methyl transferase, which adds methyl residues to the same glutamate residues. If the level of an attractant remains high, the level of phosphorylation of CheA ( and, therefore, CheY and CheB ) will remain low, the cell will swim smoothly, and the level of methylation of the MCPs will increase ( because CheB - P is not present to demethylate ). The MCPs no longer respond to the attractant when they are fully methylated ; therefore, even though the level of attractant might remain high, the level of CheA - P ( and CheB - P ) increases and the cell begins to tumble. The MCPs can be demethylated by CheB - P, and, when this happens, the receptors can once again respond to attractants. The situation is the opposite with regard to repellents : fully methylated MCPs respond best to repellents, while least - methylated MCPs respond worst to repellents. This regulation allows the bacterium to ' remember ' chemical concentrations from the recent past, a few seconds, and compare them to those it is currently experiencing, thus ' know ' whether it is traveling up or down a gradient. Although the methylation system accounts for the wide range of sensitivity that bacteria have to chemical gradients, other mechanisms are involved in increasing the absolute value of the sensitivity on a given background. Well - established examples are the ultra-sensitive response of the motor to the CheY - P signal, and the clustering of chemoreceptors. ( Eukaryotic chemotaxis This section needs additional citations for verification. Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed. ( March 2017 ) The mechanism that eukaryotic cells employ is quite different from that in bacteria ; however, sensing of chemical gradients is still a crucial step in the process. Due to their size, prokaryotes can not detect effective concentration gradients ; therefore, these cells scan and evaluate their environment by a constant swimming ( consecutive steps of straight swims and tumbles ). In contrast, the size of eukaryotic cells allows for the possibility of detecting gradients, which results in a dynamic and polarized distribution of receptors ; induction of these receptors by chemoattractants or chemorepellents results in migration towards or away from the chemotactic substance. Levels of receptors, intracellular signalling pathways and the effector mechanisms all represent diverse, eukaryotic - type components. In eukaryotic unicellular cells, amoeboid movement and cilium or the eukaryotic flagellum are the main effectors ( e.g., Amoeba or Tetrahymena ). Some eukaryotic cells of higher vertebrate origin, such as immune cells also move to where they need to be. Besides immune competent cells ( granulocyte, monocyte, lymphocyte ) a large group of cells -- considered previously to be fixed into tissues -- are also motile in special physiological ( e.g., mast cell, fibroblast, endothelial cells ) or pathological conditions ( e.g., metastases ). Chemotaxis has high significance in the early phases of embryogenesis as development of germ layers is guided by gradients of signal molecules. Motility This section does not cite any sources. Please help improve this section by adding citations to reliable sources. Unsourced material may be challenged and removed. ( March 2017 ) Unlike motility in bacterial chemotaxis, the mechanism by which eukaryotic cells physically move is unclear. There appear to be mechanisms by which an external chemotactic gradient is sensed and turned into an intracellular PIP3 gradient, which results in a gradient and the activation of a signaling pathway, culminating in the polymerisation of actin filaments. The growing distal end of actin filaments develops connections with the internal surface of the plasma membrane via different sets of peptides and results in the formation of anterior pseudopods and posterior uropods. Cilia of eukaryotic cells can also produce chemotaxis ; in this case, it is mainly a Ca - dependent induction of the microtubular system of the basal body and the beat of the 9 + 2 microtubules within cilia. The orchestrated beating of hundreds of cilia is synchronized by a submembranous system built between basal bodies. The details of the signaling pathways are still not totally clear. Chemotaxis related migratory responses This section does not cite any sources. Please help improve this section by adding citations to reliable sources. Unsourced material may be challenged and removed. ( March 2017 ) Although chemotaxis is the most frequently studied form of migration there are several other forms of locomotion in the cellular level. Chemokinesis is also induced by molecules of the liquid phase of the surrounding environment ; however, the response elicited is a not vectorial, random taxis. Neither amplitude nor frequency of motion has characteristic, directional components, as this behaviour provides more scanning of the environment than migration between two distinct points. In haptotaxis the gradient of the chemoattractant is expressed or bound on a surface, in contrast to the classical model of chemotaxis, in which the gradient develops in a soluble fluid. The most common biologically active haptotactic surface is the extracellular matrix ( ECM ) ; the presence of bound ligands is responsible for induction of transendothelial migration and angiogenesis. Necrotaxis embodies a special type of chemotaxis when the chemoattractant molecules are released from necrotic or apoptotic cells. Depending on the chemical character of released substances, necrotaxis can accumulate or repel cells, which underlines the pathophysiological significance of this phenomenon. Receptors In general, eukaryotic cells sense the presence of chemotactic stimuli through the use of 7 - transmembrane ( or serpentine ) heterotrimeric G - protein - coupled receptors, a class representing a significant portion of the genome. Some members of this gene superfamily are used in eyesight ( rhodopsins ) as well as in olfaction ( smelling ). The main classes of chemotaxis receptors are triggered by : formyl peptides - formyl peptide receptors ( FPR ), chemokines - chemokine receptors ( CCR or CXCR ), and leukotrienes - leukotriene receptors ( BLT ). However, induction of a wide set of membrane receptors ( e.g., amino acids, insulin, vasoactive peptides ) also elicit migration of the cell. Chemotactic selection While some chemotaxis receptors are expressed in the surface membrane with long - term characteristics, as they are determined genetically, others have short - term dynamics, as they are assembled ad hoc in the presence of the ligand. The diverse features of the chemotaxis receptors and ligands allows for the possibility of selecting chemotactic responder cells with a simple chemotaxis assay. By chemotactic selection, we can determine whether a still - uncharacterized molecule acts via the long - or the short - term receptor pathway. The term chemotactic selection is also used to designate a technique that separates eukaryotic or prokaryotic cells according to their chemotactic responsiveness to selector ligands. Chemotactic ligands The number of molecules capable of eliciting chemotactic responses is relatively high, and we can distinguish primary and secondary chemotactic molecules. The main groups of the primary ligands are as follows : Formyl peptides are di -, tri -, tetrapeptides of bacterial origin, formylated on the N - terminus of the peptide. They are released from bacteria in vivo or after decomposition of the cell ( a typical member of this group is the N - formylmethionyl - leucyl - phenylalanine ( abbreviated fMLF or fMLP ). Bacterial fMLF is a key component of inflammation has characteristic chemoattractant effects in neutrophil granulocytes and monocytes. The chemotactic factor ligands and receptors related to formyl peptides are summarized in the related article, Formyl peptide receptors. Complement 3a ( C3a ) and complement 5a ( C5a ) are intermediate products of the complement cascade. Their synthesis is joined to the three alternative pathways ( classical, lectin - dependent, and alternative ) of complement activation by a convertase enzyme. The main target cells of these derivatives are neutrophil granulocytes and monocytes as well. Chemokines belong to a special class of cytokines ; not only do their groups ( C, CC, CXC, CX C chemokines ) represent structurally related molecules with a special arrangement of disulfide bridges but also their target cell specificity is diverse. CC chemokines act on monocytes ( e.g., RANTES ), and CXC chemokines are neutrophil granulocyte - specific ( e.g., IL - 8 ). Investigations of the three - dimensional structures of chemokines provided evidence that a characteristic composition of beta - sheets and an alpha helix provides expression of sequences required for interaction with the chemokine receptors. Formation of dimers and their increased biological activity was demonstrated by crystallography of several chemokines, e.g. IL - 8. Metabolites of polyunsaturated fatty acids Leukotrienes are eicosanoid lipid mediators made by the metabolism of arachidonic acid by ALOX5 ( also termed 5 - lipoxygenase ). Their most prominent member with chemotactic factor activity is leukotriene B4, which elicits adhesion, chemotaxis, and aggregation of leukocytes. The chemoattractant action of LTB4 is induced via either of two G protein -- coupled receptors, BLT 1 and BLT2, which are highly expressed in cells involved in inflammation and allergy. The family of 5 - Hydroxyicosatetraenoic acid eicosanoids are arachidonic acid metabolites also formed by ALOX5. Three members of the family form naturally and have prominent chemotactic activity. These, listed in order of decreasing potency, are : 5 - oxo - eicosatetraenoic acid, 5 - oxo - 15 - hydroxy - eicosatetraenoic acid, and 5 - Hydroxyeicosatetraenoic acid. This family of agonists stimulates chemotactic responses in human eosinophils, neutrophils, and monocytes by binding to the Oxoeicosanoid receptor 1, which like the receptors for leukotriene B4, is a G protein - coupled receptor. Aside from the skin, neutrophils are the body 's first line of defense against bacterial infections. After leaving nearby blood vessels, these cells recognize chemicals produced by bacteria in a cut or scratch and migrate `` toward the smell ''. 5 - hydroxyeicosatrieonic acid and 5 - oxoeicosatrienoic acid are metabolites of Mead acid ( 5Z, 8Z, 11Z - eicosatrirenoid acid ) ; they stimulate leukocyte chemotaxis through the oxoeicosanoid receptor 1 with 5 - oxoeicosatrienoic acid being as potent as its arachidonic acid - derived analog, 5 - oxo - eicosatetraenoic acid, in stimulating human blood eosinophil and neutrophil chemotaxis. 12 - Hydroxyeicosatetraenoic acid is an eicosanoid metabolite of arachidonic acid made by ALOX12 which stimulates leukocyte chemotaxis through the leukotriene B4 receptor, BLT2. Prostaglandin D2 is an eicosanoid metabolite of arachidononic acid made by cyclooxygenase 1 or cyclooxygenase 2 that stimulates chemotaxis through the Prostaglandin DP2 receptor. It elicits chemotactic responses in eosinophils, basophils, and T helper cells of the Th2 subtype. 12 - Hydroxyheptadecatrienoic acid is a non-eicosanoid metabolite of arachidonic acid made by cyclooxygenase 1 or cyclooxygenase 2 that stimulates leukocyte chemataxis though the leukotriene B4 receptor, BLT2. 15 - oxo - eicosatetraenoic acid is an eicosanoid metabolite of arachidonic acid made my ALOX15 ; it has weak chemotactic activity for human monocytes ( sees 15 - Hydroxyeicosatetraenoic acid # 15 - oxo - ETE ). The receptor or other mechanism by which this metabolite stimulates chemotaxis has not been elucidated. Chemotactic range fitting This section has multiple issues. Please help improve it or discuss these issues on the talk page. ( Learn how and when to remove these template messages ) This section does not cite any sources. Please help improve this section by adding citations to reliable sources. Unsourced material may be challenged and removed. ( March 2017 ) This article possibly contains original research. Please improve it by verifying the claims made and adding inline citations. Statements consisting only of original research should be removed. ( March 2017 ) Chemotactic responses elicited by the ligand - receptor interactions are, in general, distinguished upon the optimal effective concentration ( s ) of the ligand. Nevertheless, correlation of the amplitude elicited and ratio of the responder cells compared to the total number are also characteristic features of the chemotactic signaling. Investigations of ligand families ( e.g., amino acids or oligo peptides ) proved that there is a fitting of ranges ( amplitudes ; number of responder cells ) and chemotactic activities : Chemoattractant moiety is accompanied by wide ranges, whereas chemorepellent character by narrow ranges. Clinical significance This section does not cite any sources. Please help improve this section by adding citations to reliable sources. Unsourced material may be challenged and removed. ( March 2017 ) A changed migratory potential of cells has relatively high importance in the development of several clinical symptoms and syndromes. Altered chemotactic activity of extracellular ( e.g., Escherichia coli ) or intracellular ( e.g., Listeria monocytogenes ) pathogens itself represents a significant clinical target. Modification of endogenous chemotactic ability of these microorganisms by pharmaceutical agents can decrease or inhibit the ratio of infections or spreading of infectious diseases. Apart from infections, there are some other diseases wherein impaired chemotaxis is the primary etiological factor, as in Chédiak -- Higashi syndrome, where giant intracellular vesicles inhibit normal migration of cells. Chemotaxis in diseases Type of disease Chemotaxis increased Chemotaxis decreased Infections inflammations AIDS, Brucellosis Chemotaxis results the disease -- Chédiak -- Higashi syndrome, Kartagener syndrome Chemotaxis is affected atherosclerosis, arthritis, periodontitis, psoriasis, reperfusion injury, metastatic tumors multiple sclerosis, Hodgkin disease, male infertility Intoxications asbestos, benzpyrene Hg and Cr salts, ozone Mathematical models This section needs additional citations for verification. Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed. ( March 2017 ) Several mathematical models of chemotaxis were developed depending on the type of migration ( e.g., basic differences of bacterial swimming, movement of unicellular eukaryotes with cilia / flagellum and amoeboid migration ) physico - chemical characteristics of the chemicals ( e.g., diffusion ) working as ligands biological characteristics of the ligands ( attractant, neutral, and repellent molecules ) assay systems applied to evaluate chemotaxis ( see incubation times, development, and stability of concentration gradients ) other environmental effects possessing direct or indirect influence on the migration ( lighting, temperature, magnetic fields, etc. ) Although interactions of the factors listed above make the behavior of the solutions of mathematical models of chemotaxis rather complex, it is possible to describe the basic phenomenon of chemotaxis - driven motion in a straightforward way. Indeed, let us denote with φ ( displaystyle \varphi ) the spatially non-uniform concentration of the chemo - attractant and with ∇ φ ( displaystyle \nabla \varphi ) its gradient. Then the chemotactic cellular flow ( also called current ) J ( displaystyle J ) that is generated by the chemotaxis is linked to the above gradient by the law : J = χ C ∇ φ ( displaystyle J = chi C \nabla \varphi ), where C ( displaystyle C ) is the spatial density of the cells and χ ( displaystyle chi ) is the so - called ' Chemotactic coefficient '. However, note that in many cases χ ( displaystyle chi ) is not constant : It is, instead, a decreasing function of the concentration of the chemo - attractant φ ( displaystyle \varphi ) : χ ( φ ) ( displaystyle chi ( \varphi ) ). Spatial ecology of soil microorganisms is a function of their chemotactic sensitivities towards substrate and fellow organisms. The chemotactic behavior of the bacteria was proven to lead to non-trivial population patterns even in the absence of environmental heterogeneities. The presence of structural pore scale heterogeneities has an extra impact on the emerging bacterial patterns. Measurement of chemotaxis This section does not cite any sources. Please help improve this section by adding citations to reliable sources. Unsourced material may be challenged and removed. ( March 2017 ) Main article : Chemotaxis assay A wide range of techniques is available to evaluate chemotactic activity of cells or the chemoattractant and chemorepellent character of ligands. The basic requirements of the measurement are as follows : concentration gradients can develop relatively quickly and persist for a long time in the system chemotactic and chemokinetic activities are distinguished migration of cells is free toward and away on the axis of the concentration gradient detected responses are the results of active migration of cells Despite the fact that an ideal chemotaxis assay is still not available, there are several protocols and pieces of equipment that offer good correspondence with the conditions described above. The most commonly used are summarised in the table below : Type of assay Agar - plate assays Two - chamber assays Others Examples PP - chamber Boyden chamber Zigmond chamber Dunn chambers Multi-well chambers Capillary techniques T - maze technique Opalescence technique Orientation assays Artificial chemotactic systems Chemical robots that use artificial chemotaxis to navigate autonomously have been designed. Applications include targeted delivery of drugs in the body. See also McCutcheon index Tropism Durotaxis Haptotaxis Mechanotaxis Plithotaxis References Jump up ^ Chisholm, Hugh, ed. ( 1911 ). `` Chemotaxis ''. Encyclopædia Britannica. 6 ( 11th ed. ). Cambridge University Press. p. 77. Jump up ^ Chemotaxis Lecture. Uploaded in 2007. available at : http://www.rpgroup.caltech.edu/courses/aph161/2007/lectures/ChemotaxisLecture.pdf ( Last inspected : 15 / 04 / 17 ) Jump up ^ Élie Metchnikoff ``. Encyclopædia Britannica. Encyclopædia Britannica, Inc. Jump up ^ Keller - Segel Models for Chemotaxis. 2012. available at : http://www.isn.ucsd.edu/courses/Beng221/problems/2012/BENG221_Project%20-%20Roberts%20Chung%20Yu%20Li.pdf ( Last inspected by April 2017 ) Jump up ^ Snyderman R, Gewurz H, Mergenhagen SE ( 1968 ). `` Interactions of the Complement System with Endotoxic Lipopolysaccharide. Generation of a Factor Chemotactic for Polymorphonuclear L eukocytes ''. J. Exp. Med. 128 ( 2 ; August ) : 259 -- 275. CS1 maint : Multiple names : authors list ( link ) Jump up ^ Adler J & Tso, W-W ( 1974 ). `` Decision - Making in Bacteria : Chemotactic Response of Escherichia Coli to Conflicting Stimuli ''. Science. 184 ( 4143 ) : 1292 -- 1294. Bibcode : 1974Sci... 184.1292 A. PMID 4598187. doi : 10.1126 / science. 184.4143. 1292. Jump up ^ Macnab RM, Koshland DE ( September 1972 ). `` The gradient - sensing mechanism in bacterial chemotaxis ''. Proc. Natl. Acad. Sci. U.S.A. 69 ( 9 ) : 2509 -- 12. Bibcode : 1972PNAS... 69.2509 M. PMC 426976. PMID 4560688. doi : 10.1073 / pnas. 69.9. 2509. Jump up ^ Berg, Howard C. ( 2003 ). E. coli in motion. New York, NY : Springer. ISBN 0387008888. Jump up ^ ToxCafe ( 2 June 2011 ). `` Chemotaxis ''. Retrieved March 23, 2017 -- via YouTube. Jump up ^ Mello BA & Tu Y ( 2007 ). `` Effects of Adaptation in Maintaining High Sensitivity Over a Wide Range of Backgrounds for Escherichia coli Chemotaxis ''. 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PMID 9702410. doi : 10.1006 / cyto. 1997.0328. ^ Jump up to : Powell WS, Rokach J ( 2015 ). `` Biosynthesis, biological effects, and receptors of hydroxyeicosatetraenoic acids ( HETEs ) and oxoeicosatetraenoic acids ( oxo - ETEs ) derived from arachidonic acid ''. Biochimica et Biophysica Acta. 1851 ( 4 ) : 340 -- 55. PMID 25449650. doi : 10.1016 / j. bbalip. 2014.10. 008. Retrieved March 23, 2017. Jump up ^ Powell WS, Rokach J ( 2013 ). `` The eosinophil chemoattractant 5 - oxo - ETE and the OXE receptor ''. Prog. Lipid Res. 52 ( 4 ; October ) : 651 -- 65. doi : 10.1016 / j. plipres. 2013.09. 001. Retrieved March 23, 2017. Jump up ^ Matsuoka T, Narumiya S ( 2007 ). `` Prostaglandin receptor signaling in disease ''. The Scientific World Journal. 7 : 1329 -- 47. PMID 17767353. doi : 10.1100 / tsw. 2007.182. Jump up ^ Yokomizo T ( 2015 ). `` Two distinct leukotriene B4 receptors, BLT1 and BLT2 ''. Journal of Biochemistry. 157 ( 2 ) : 65 -- 71. PMID 25480980. doi : 10.1093 / jb / mvu078. Jump up ^ Sozzani, S ; Zhou, D ; Locati, M ; Bernasconi, S ; Luini, W ; Mantovani, A ; O'Flaherty, J.T. ( 1996 ). `` Stimulating properties of 5 - oxo - eicosanoids for human monocytes : Synergism with monocyte chemotactic protein - 1 and - 3 ''. Journal of Immunology. 157 ( 10 ) : 4664 -- 71. PMID 8906847. Jump up ^ Gharasoo, Mehdi ; Centler, Florian ; Fetzer, Ingo ; Thullner, Martin ( 2014 ). `` How the chemotactic characteristics of bacteria can determine their population patterns ''. Soil Biology and Biochemistry. 69 : 346. doi : 10.1016 / j. soilbio. 2013.11. 019. Jump up ^ Lagzi, István ( 2013 ). `` Chemical Robotics -- Chemotactic Drug Carriers ''. Central European Journal of Medicine. 8 ( 4 ) : 377 -- 382. doi : 10.2478 / s11536 - 012 - 0130 - 9. Further reading Alberts, Bruce ; Johnson, Alexander ; Lewis, Julian ; Walter, Peter ; Raff, Martin C. ( 2002 ). `` Bacterial Chemotaxis Depends on a Two - Component Signaling Pathway Activated by Histidine - Kinase - associated Receptors ''. Molecular Biology of the Cell. Taylor & Francis Group. ISBN 9780815340690. Retrieved 18 September 2017. Bagorda, A. ; Parent, C.A. ( 6 August 2008 ). `` Eukaryotic chemotaxis at a glance ''. Journal of Cell Science. 121 ( 16 ) : 2621 -- 2624. doi : 10.1242 / jcs. 018077. Berg, Howard C. ( 1993 ). Random walks in biology ( Expanded, rev. ed. ). Princeton, NJ : Princeton Univ. Press. ISBN 9780691000640. Berg, Howard C. ( 2003 ). E. coli in motion. New York : Springer. ISBN 9780387008882. Dusenbery, David B. ( 2009 ). Living at micro scale : the unexpected physics of being small. Cambridge, Mass. : Harvard University Press. ISBN 9780674031166. Eisenbach, Michael ( 2004 ). Lengeler, Joseph W., ed. Chemotaxis. London : Imperial College Press. ISBN 9781860944130. Eisenbach, Michael ( December 2011 ). `` Bacterial Chemotaxis ''. eLS. doi : 10.1002 / 9780470015902. a0001251. pub3. Hazelbauer, Gerald L. ( 13 October 2012 ). `` Bacterial Chemotaxis : The Early Years of Molecular Studies ''. Annual Review of Microbiology. 66 ( 1 ) : 285 -- 303. doi : 10.1146 / annurev - micro-092611 - 150120. Jin, Tian ; Hereld, Dale ( 2016 ). Chemotaxis : Methods and Protocols. Humana Press. ISBN 9781493934805. L.D., Miller ; M.H., Russell ; G., Alexandre ( 2009 ). `` Diversity in bacterial chemotactic responses and niche adaptation. ''. Advances in Applied Microbiology. 66 : 53 -- 75. doi : 10.1016 / S0065 - 2164 ( 08 ) 00803 - 4. Rao CV, Kirby JR & Arkin AP ( 2004 ). `` Design and Diversity in Bacterial Chemotaxis : A Comparative Study in Escherichia coli and Bacillus subtilis ''. PLoS Biol. 2 ( 2 ) : e49. doi : 10.1371 / journal. pbio. 0020049. Williams, Ann H. ( 20 December 2010 ). `` Chemotaxis on the Move -- Active Learning Teaching Tool ''. Journal of Microbiology & Biology Education. 11 ( 2 ) : 177 -- 178. doi : 10.1128 / jmbe. v11i2. 216. External links Wikibooks has more on the topic of : Chemotaxis Wikimedia Commons has media related to Chemotaxis. Chemotaxis Neutrophil Chemotaxis Cell Migration Gateway Downloadable Matlab chemotaxis simulator Bacterial Chemotaxis Interactive Simulator ( web - app ) Retrieved from `` https://en.wikipedia.org/w/index.php?title=Chemotaxis&oldid=804739701 '' Categories : Motile cells Perception Taxes ( biology ) Transmembrane receptors Hidden categories : Wikipedia articles incorporating a citation from the 1911 Encyclopaedia Britannica with Wikisource reference CS1 maint : Multiple names : authors list All pages needing factual verification Wikipedia articles needing factual verification from March 2017 Wikipedia articles needing page number citations from March 2017 Articles needing additional references from September 2015 All articles needing additional references Articles lacking reliable references from March 2017 All articles lacking reliable references Articles needing expert attention from March 2017 All articles needing expert attention Physiology articles needing expert attention Molecular biology articles needing expert attention Articles covered by WikiProject Wikify from March 2017 Articles needing footnote reformatting All articles covered by WikiProject Wikify Articles needing additional references from March 2017 Articles with specifically marked weasel - worded phrases from April 2017 Articles to be expanded from March 2017 All articles to be expanded All articles with unsourced statements Articles with unsourced statements from March 2017 All articles with specifically marked weasel - worded phrases Articles with specifically marked weasel - worded phrases from March 2017 Articles that may contain original research from March 2017 All articles that may contain original research Talk Contents About Wikipedia Català Čeština Dansk Deutsch Ελληνικά Español Esperanto Euskara فارسی Français Gaeilge Italiano עברית Қазақша Magyar Nederlands 日本 語 Polski Português Română Русский Slovenčina Српски / srpski Srpskohrvatski / српскохрватски Svenska தமிழ் Українська اردو 中文 Edit links This page was last edited on 10 October 2017, at 21 : 05. 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Question: the bacterium e. coli swims in a straight line called a
Answer: Chemotaxis