A taxis (from
Ancient Greekτάξις (táxis) 'arrangement, order';[1]
pl.: taxes/ˈtæksiːz/)[2][3][4] is the
movement of an
organism in response to a
stimulus such as light or the presence of food. Taxes are
innate behavioural responses. A taxis differs from a
tropism (turning response, often growth towards or away from a stimulus) in that in the case of taxis, the organism has
motility and demonstrates guided movement towards or away from the stimulus source.[5][6] It is sometimes distinguished from a
kinesis, a non-directional change in activity in response to a stimulus.
Classification
Taxes are classified based on the type of stimulus, and on whether the organism's response is to move towards or away from the stimulus. If the organism moves towards the stimulus the taxis are positive, while if it moves away the taxis are negative. For example, flagellate
protozoans of the genus Euglena move towards a light source. This reaction or behavior is called positive phototaxis since phototaxis refers to a response to light and the organism is moving towards the stimulus.
Terminology derived from type of stimulus
Many types of taxis have been identified, including:
Depending on the type of
sensory organs present, a taxis can be classified as a klinotaxis, where an organism continuously samples the environment to determine the direction of a stimulus; a tropotaxis, where bilateral sense organs are used to determine the stimulus direction; and a telotaxis, where a single organ suffices to establish the orientation of the stimulus.
Terminology derived from taxis direction
There are five types of taxes based on the movement of organisms.
Klinotaxis occurs in organisms with
receptor cells but not paired receptor organs. The cells for reception may be located all over the body, but often towards the anterior side. The organism detects the
stimuli by turning its head sideways and comparing the intensity of the stimulus. Their direction of movement is then based on the stronger stimulus, either moving toward a desirable stimulus or away from an undesired one.[7] When the intensity of stimuli is balanced equally from all sides, the organism moves in a straight line. The movement of
blowfly and
butterflylarvae clearly demonstrates klinotaxis.
Tropotaxis is displayed by
organisms with paired receptor cells, comparing the strength of the signals and turning toward the strongest signal.[7] The movement of
grayling butterflies and
fish lice clearly demonstrates tropotaxis.
Telotaxis also requires paired receptors. The movement occurs along the direction where the intensity of the stimuli is stronger. Telotaxis is clearly seen in the movement of
bees when they leave their
hive to look for food. They balance the stimuli from the sun as well as from
flowers but land on the flower whose stimulus is most intense for them.
Menotaxis describes organisms' maintenance of a constant
angular orientation. A clear demonstration is shown by bees returning to their hive at night and the movement of ants with respect to the sun.
Mnemotaxis is the use of
memory to follow trails that organisms have left when travelling to or from their home.
Examples
Aerotaxis is the response of an organism to variation in oxygen concentration, and is mainly found in aerobic bacteria.[8]
Anemotaxis is the response of an organism to wind. Many insects show a positive anemotactic response (turning/flying into the wind) upon exposure to an airborne stimulus cue from a food source or pheromones.[7] Cross-wind anemotactic search is exhibited by some olfactory animals in the absence of a target odor including moths, albatrosses, and polar bears.[9][10][11] Rats have specialized supra-orbital whiskers that detect wind and cause anemotactic turning.[12]
Chemotaxis is a response elicited by chemicals: that is, a response to a chemical concentration gradient.[8][7][13] For example, chemotaxis in response to a sugar gradient has been observed in motile bacteria such as
E. coli.[14] Chemotaxis also occurs in the
antherozoids of
liverworts,
ferns, and
mosses in response to chemicals secreted by the
archegonia.[8] Unicellular (e.g. protozoa) or multicellular (e.g. worms) organisms are targets of chemotactic substances. A concentration gradient of chemicals developed in a fluid phase guides the vectorial movement of responder cells or organisms. Inducers of locomotion towards increasing steps of concentrations are considered as
chemoattractants, while
chemorepellents result moving off the chemical. Chemotaxis is described in
prokaryotic and
eukaryotic cells, but signalling mechanisms (receptors, intracellular signaling) and effectors are significantly different.
Durotaxis is the directional movement of a cell along a stiffness gradient.
Electrotaxis (or galvanotaxis) is the directional movement of motile cells along the vector of an
electric field. It has been suggested that by detecting and orienting themselves toward the electric fields, cells can move towards damages or wounds to repair them. It also is suggested that such a movement may contribute to directional growth of cells and tissues during development and regeneration. This notion is based on the existence of measurable electric fields that naturally occur during wound healing, development and regeneration; and cells in cultures respond to applied electric fields by directional cell migration – electrotaxis / galvanotaxis.
Energy taxis is the orientation of bacteria towards conditions of optimal metabolic activity by sensing the internal energetic conditions of cell. Therefore, in contrast to chemotaxis (taxis towards or away from a specific extracellular compound), energy taxis responds on an intracellular stimulus (e.g.
proton motive force, activity of
NDH- 1) and requires metabolic activity.[15]
Magnetotaxis is, strictly speaking, the ability to sense a
magnetic field and coordinate movement in response. However, the term is commonly applied to bacteria that contain magnets and are physically rotated by the force of
Earth's magnetic field. In this case, the "behaviour" has nothing to do with sensation and the bacteria are more accurately described as "magnetic bacteria".[19]
Pharotaxis is the movement to a specific location in response to learned or conditioned stimuli, or navigation by means of landmarks.[20][21]
Phonotaxis is the movement of an organism in response to
sound.
Phototaxis is the movement of an organism in response to
light: that is, the response to variation in light intensity and direction.[8][22] Negative phototaxis, or movement away from a light source, is demonstrated in some insects, such as cockroaches.[8] Positive phototaxis, or movement towards a light source, is advantageous for phototrophic organisms as they can orient themselves most efficiently to receive light for
photosynthesis. Many
phytoflagellates, e.g. Euglena, and the
chloroplasts of higher plants positively phototactic, moving towards a light source.[8] Two types of positive phototaxis are observed in prokaryotes: scotophobotaxis is observable as the movement of a bacterium out of the area illuminated by a microscope, when entering darkness signals the cell to reverse direction and reenter the light; a second type of positive phototaxis is true phototaxis, which is a directed movement up a gradient to an increasing amount of light. There is a different classification to orientation towards dark areas called scototaxis.
Rheotaxis is a response to a
current in a fluid. Positive rheotaxis is shown by fish turning to face against the current. In a flowing stream, this behaviour leads them to hold their position in a stream rather than being swept downstream. Some fish will exhibit negative rheotaxis where they will avoid currents.
Thermotaxis is a migration along a gradient of temperature. Some
slime molds and small
nematodes can migrate along amazingly small temperature gradients of less than 0.1 °C/cm.[23] They apparently use this behaviour to move to an optimal level in soil.[24][25]
Thigmotaxis is the response of an organism to physical contact or to the proximity of a physical discontinuity in the environment (e.g. rats preferring to swim near the edge of a water maze).
Codling moth larvae are believed to use thigmotactic sense to locate fruits to feed on.[26] Mice and rats, when inhabiting human-made structures, tend to stick close to vertical surfaces; this primarily manifests as running along the floor/wall juncture. Whiskers (
vibrissae) are often used to detect the presence of a wall or surface in the absence of sufficient light in rodents and felines to aid in thigmotaxis.
^Blass, E.M (1987). "Opioids, sweets and a mechanism for positive affect: Broad motivational implications". In Dobbing, J (ed.). Sweetness. London: Springer-Verlag. pp. 115–124.
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^C. F. Adams & A. J. Paul (1999). "Phototaxis and geotaxis of light-adapted zoeae of the golden king crab Lithodes aequispinus (Anomura: Lithodidae) in the laboratory". Journal of Crustacean Biology. 19 (1): 106–110.
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^Menzel, Randolf (1979). "Spectral Sensitivity and Color Vision in Invertebrates". In H. Autrum (ed.). Comparative Physiology and Evolution of Vision in Invertebrates- A: Invertebrate Photoreceptors. Handbook of Sensory Physiology. Vol. VII/6A. New York: Springer-Verlag. pp. 503–580. See section D: Wavelength–Specific Behavior and Color Vision.
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^Dusenbery, David B. (1992). Sensory Ecology, p.114. W.H. Freeman, New York.
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^Dusenbery, D.B. Behavioral Ecology and Sociobiology, 22:219–223 (1988). "Avoided temperature leads to the surface:…"
^Dusenbery, D.B. Biological Cybernetics, 60:431–437 (1989). "A simple animal can use a complex stimulus patter to find a location:…"