Chordotonal organ

Chordotonal organs are stretch receptor organs in insects and other arthropods.[1] They are located at most joints[2] and are made up of clusters of scolopidia that either directly or indirectly connect two joints and sense their movements relative to one another. They can have both extero- and proprioceptive functions, for example sensing auditory stimuli or leg movement.[3] The femoral chordotonal organ is thought to be functionally homologous to muscle spindles.[4] The word was coined by Vitus Graber in 1882 though he interpreted them as being stretched between two points like a string, sensing vibrations through resonance.[5]

Structure
Diagram of the primary components of a chordotonal organ scolopidium

Chordotonal organs can be composed of a single scolopidium with only a single sensory, bipolar neuron (such as the tympanal ear of a notodontid moth), or up to several thousand scolopidia, each equipped with up to four sensory neurons (as in the mosquito Johnston's organ).[6] The bipolar sensory neurons each have an apical dendritic structure with a cilium densely packed with microtubules and surrounded by two specialized cells, the scolopale cell and the attachment (cap) cell, plus a glial cell.[2] Mechanically gated ion channels are located distal to the ciliary dilation, a characteristic part of the upper dendiritic cilium. The cavity between the scolopale cell and the sensory neuron is filled with a specialized receptor lymph similar to the endolymph that surrounds the mechanosensory hair bundles of cochlear hair cells (high in potassium and low in sodium).[6] The dendritic cilia can have one of two major forms: in the mononematic form, the major connection between the attachment site and the cilium is a microtubule-rich attachment cell. The electron-dense extracellular material is small and localized mainly to the junction between the cilia and the attachment cell. The femoral chordotonal organ is mononematic. In contrast, in the amphinematic form, the extracellular material of the cap forms a dense, tubular sheath that surrounds the sensory cilium and extends all the way to the cuticle at the attachment site. In this form, the attachment cell contains both microtubules and actin-rich scolopale rods similar to those present in the scolopale cell. The Johnston's organ is an example of an amphinematic chordotonal organ. The functional significance of the morphological differences of the two forms is unknown, but may confer different viscoelastic properties on the sensory units.[7]

Functional diversity

In a chordotonal organ, individual sensory neurons can respond to different types of mechanosensory stimuli (for example, sound vs gravity), and those that respond to a particular stimulus can have different tuning properties (for example, tuned to different position of a joint)[2]. One way to generate these functional diversity is by having sensory neurons with different types of mechanosensory channels or intrinsic properties. For example, in Johnston's organ of Drosophila melanogaster, sensory neurons that detect sound may express nompC, an ion channel that belongs to the transient receptor potential (TRP) superfamily, while those that detect gravity may express another member of the TRP channel, painless[8]. Another way to generate functional diversity is by having sensory neurons that are attached to the joint through different types of connections. For example, in the femoral chordotonal organ of the locust, the ligament in which sensory neurons are embedded is divided into several strands that are sequentially pulled as the joint is flexed, providing a mechanism for differential activation of the sensory neurons at different position of the joint[9].

Major chordotonal organs

Femoral chordotonal organ

The femoral chordotonal organ is located in the femur of the insect leg and it detects position, speed, acceleration, and vibration of the tibia relative to the femur.[10][11][12][13][14] In Drosophila melanogaster, where it is possible to systematically analyze neuronal populations using genetic tools, the sensory neurons of the femoral chordotonal organ can be separated into at least 3 functionally and genetically distinct populations: the club, claw, and hook.[14] The club neurons encode bi-directional movements and vibrations of the tibia, the claw neurons encode position of the tibia, and the hook neurons encode directional movements of the tibia. [14] Information encoded by the femoral chordotonal organ is thought to be used during behaviors that require precise control of leg movements, such as walking [15]and target reaching[16].

In the femoral chordotonal organ, the scolopidia are organized into groups called scoloparia. Scoloparia may be functionally distinct from one another, with separate scoloparia containing vibration-sensitive or position-sensitive sensory neurons.[17][18] D. melanogaster has three scoloparia.[19]

Johnston's organ

The chordotonal organ at the base of the antennae is called the Johnston's organ, and it detects movement of the antennae. Johnston's organ occurs in nearly all orders of insects.[2] In Drosophila melanogaster, one population of Johnston's organ neurons has been shown to mediate the sensation of sound, whereas another population detects wind and gravity. In several species of Diptera, the Johnston's organ is sexually dimorphic. Males possess both greater numbers, greater diversity, and a more highly organized distribution of scolopidia.[3] Some species of mosquitoes may possess as many as several thousand scolopidia.[6] Males of these species likely use the Johnston's organ to identify potential mates.

Janet's organ

In addition to the Johnston's organ, antennae of Hymenoptera possess a second chordotonal organ, the Janet's organ, which detects flexion of the antennal joints, somewhat like the femoral chordotonal organ.[3]

Subgenual organ

The subgenual organ is found in all insects except Diptera and Coleoptera. It is located in the proximal part of the tibia and detects high-frequency acoustic vibrations transmitted through the substrate as well as sound through air.[20]

Tympanal organ

Tympanal organs are specialized hearing organs that have evolved in at least seven different orders of insects. They consist of a tympanal membrane backed by an air-filled space and are innervated by a chordotonal organ. Tympanal organs detect air-borne vibrations and are used to detect predators, prey, and potential mates and rivals. They can be found in a variety of locations on the body, including the abdomen, wing base, metathorax, and ventral prosternum.[21]

Wing and halteres

There is a chordotonal organ located at the base of the wings in many insect orders, and, in Dipterans, there are also two chordotonal organs found at the base of the haltere. Their function is currently not well understood. In lacewings, a tympanal organ is located in the radius vein of the forewing and is thought to monitor ultrasound.[2]

See also

References
  1. Christensen TA (December 2004). Methods in Insect Sensory Neuroscience. CRC Press. ISBN 978-1-4200-3942-9.
  2. Field LH, Matheson T (January 1998). "Chordotonal organs of insects" (PDF). Advances in Insect Physiology. 27: 1–228. doi:10.1016/S0065-2806(08)60013-2. ISBN 9780120242276.
  3. Krishnan A, Sane SP (2015), "Antennal Mechanosensors and Their Evolutionary Antecedents", Advances in Insect Physiology, Elsevier, pp. 59–99, doi:10.1016/bs.aiip.2015.06.003, ISBN 978-0-12-802586-4
  4. Tuthill JC, Azim E (March 2018). "Proprioception". Current Biology. 28 (5): R194–R203. doi:10.1016/j.cub.2018.01.064. PMID 29510103.
  5. Dethier, V.G. (1963). The Physiology of Insect Senses. London: Methuen & Co. Ltd. p. 21.
  6. Kavlie RG, Albert JT (May 2013). "Chordotonal organs". Current Biology. 23 (9): R334-5. doi:10.1016/j.cub.2013.03.048. PMID 23660347.
  7. Eberl DF, Boekhoff-Falk G (2007). "Development of Johnston's organ in Drosophila". The International Journal of Developmental Biology. 51 (6–7): 679–87. doi:10.1387/ijdb.072364de. PMC 3417114. PMID 17891726.
  8. Sun, Yishan; Liu, Lei; Ben-Shahar, Yehuda; Jacobs, Julie S.; Eberl, Daniel F.; Welsh, Michael J. TRPA channels distinguish gravity sensing from hearing in Johnston's organ. National Academy of Sciences. OCLC 678802832.
  9. Field, Laurence H. (1991-02-01). "Mechanism for range fractionation in chordotonal organs of Locusta migratoria (L) and Valanga sp. (Orthoptera : Acrididae)". International Journal of Insect Morphology and Embryology. 20 (1): 25–39. doi:10.1016/0020-7322(91)90025-5. ISSN 0020-7322.
  10. Hofmann T, Koch UT, Bassler U (1985). "Physiology of the femoral chordotonal organ in the stick insect Cuniculina impigra". J. Exp. Biol. 114: 207-223.
  11. Hofmann, T (1985). "Acceleration receptors in the femoral chordotonal organ in the stick insect Cuniculina impigra". J. Exp. Biol. 114: 225-237.
  12. Zill, S (1985). "Plasticity and proprioception in the insects. I. Responses and cellular properties of individual receptors of the locust metathoracic femoral chordotonal organ". J. Exp. Biol. 116: 435-461.
  13. Matheson, T (1990). "Responses and locations of neurones in the locust metathoracic femoral chordotonal organ". J. Comp. Physiol. A. 166: 915-927.
  14. Mamiya A, Gurung P, Tuthill JC (November 2018). "Neural Coding of Leg Proprioception in Drosophila". Neuron. 100 (3): 636–650.e6. doi:10.1016/j.neuron.2018.09.009. PMC 6481666. PMID 30293823.
  15. Bassler, U (1988). "Functional principles of pattern generation for walking movements of stick insect forelegs: the role of the femoral chordotonal organ afferences". J. Exp. Biol. 136: 125-147.
  16. Page KL, Matheson T (2009). "Functional recovery of aimed scratching movements after a graded proprioceptive manipulation". J. Neurosci. 29: 3897-3907.
  17. Burns, M.D. (1974). "Structure and physiology of the locust femoral chordotonal organ". J. Insect. Physiol. 20: 1319-1339.
  18. Field, L.H.; Pfluger, H.J. (1989). "The femoral chordotonal organ: a bifunctional orthopteran (Locusta migratoria) sense organ". Comp. Biochem. Physiol. 93A: 729-743.
  19. Shanbhag, S.R.; Singh, K; Singh, R. N. (1992). "Ultrastructure of the femoral chordotonal organs and their novel synaptic organization in the legs of Drosophila melanogaster Melgen (Diptera: Drosophilidae)". Int. J. Insect Morphol. Embryol. 21: 311-322.
  20. Shaw S (August 1994). "Detection of Airborne Sound by a Cockroach 'Vibration Detector': A Possible Missing Link in Insect Auditory Evolution". The Journal of Experimental Biology. 193 (1): 13–47. PMID 9317246.
  21. Hoy RR, Robert D (January 1996). "Tympanal hearing in insects". Annual Review of Entomology. 41 (1): 433–50. doi:10.1146/annurev.en.41.010196.002245. PMID 15012336.

Further reading
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