Subgenual organ - Wikipedia

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Subgenual organ
From Wikipedia, the free encyclopedia

The subgenual organ is an organ in insects that is involved in the

perception of sound. The name (Latin sub-below and genus-knee)
refers to the location of the organ just below the knee in the tibia of
all legs in most insects. It is associated with sensing ground
movement (less than 1 nm of displacement can be enough) and
sometimes also of sound. Crickets and katydids have particularly
well-developed subgenual organs which are associated with a
tympanal organ on the tibia but probably don't share the same
function. Lepidoptera and Hymenoptera usually have less well
The neuroanatomy of the Subgenual
developed subgenual organs, in the former they are suspended on the
organ, a sensory organ in
subgenual nerve, in the latter cone shaped. In Orthoptera and
Troglophilus neglectus. SGO is the
Blattodea it often lies in the neighbourhood of other sensory organs
Subgenual organ, pIO is the proximal
containing scolopidia and is then called the SGO complex. Diptera
intermediary organ and dIO is the
and Coleoptera lack such an organ completely, Panorpa has only one
distal intermediary organ
sensory neuron, while some parasitoid wasps have as much as 400
scolopidia in their organs. In some insects, the organ is split in two
parts. In Periplaneta cockroaches, a sensitivity down to 2 nm of displacement has been determined.[1][2]
[3][4][5] The subgenual organ is particularly important in parasitoid wasps, where it is a major way of finding
suitable target animals in substrates.[6] The sensitivity of the organ varies from species to species, in
Orthoptera, Hymenoptera and Lepidoptera it is between one and more kHz, in Hemiptera only a few
hundred Hz.[1]

Development
In Teleogryllus commodus, each larval stage forms one scolopidium that contributes to the organogenesis,
while in the bushcricket Ephippiger ephippiger all scolopidia (22-24 in total) are already present in the first
larval stage. In the latter, the organ increases in size proportionally to the growth of the limb containing it
and has the shape of a fan.[7] In the cricket Teleofryllus commodus all scolopidia are formed by the third
larval stage and the organ has already its final shape by the time of egg hatching.[8] A bilobar structure in the
locust embry forms the precursor of the subgenual organ.[9] Axon growth from the subgenual organ in
grasshoppers is contingent on semaphorin I.[10]

Anatomy in specific species

In the honeybee Apis mellifera, the sensing by the subgenual organ is directed by the inertia of the
haemolymph; it causes a differential movement of the organ swimming in the haemolymph with respect to
the rest of the limb. More than 39 scolopidia, sensory cells, are involved in sensing the movement of the
haemolymph between the cuticle and two tracheae. The functionality is similar to the vestibular system of
vertebrates.[11] The bee organ is cone shaped branching out from its nerve and almost obstructs haemolymph
flow through the limb.[12]
In the carpenter ant Camponotus ligniperda has the form of a deformed sphere. On one end attachment cells
connect it to the cuticle; on the other it is innervated by the tibial nerve. The organ has the shape of a cavity
surrounded with a monocellular membrane that is heavily folded on the inside. Sensilla extend into the
cavity, each containing one neuron with associated dendrites, cilia and glial cells within a lymphatic cavity
that is connected to the cavity of the subgenual organ.[13]

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In the termite Zootermopsis angusticollis and the cockroach Periplaneta americana, the vibration is
perceived after about 10-20 milliseconds and stops being perceived after one or two seconds. There are two
types of cells with different spatial orientation in the organ; possibly, oscillation causes the cells to shift with
respect to each other and generate a signal.[14] Some early research claimed that the sensitivity of the
Periplaneta subgenual organ might be far higher than the threshold of about one atom diametre determined
for cochlear cells; newer investigation indicated that such a sensitivity may have been the result of artifacts,
with the actual sensitivity being comparable to the cochlea.[15]
In cockroaches Blaberus discoidalis and Blattella germanica, the organ has the shape of a fan that is placed
across the limb. Much of its volume is filled with discoidal cells that serve accessory purposes; they are
placed between an epidermal cell layer attached to the cuticle and connective tissue. Sensory structures
called chordotonal sensilla are involved in the perception of movement proper and contain a neuron per
sensillum, about 40-50 in total. This neuron has a single dendrite and several cilia extend from it.[16]
In Chrysoperla carnea, the green lacewing, the organ is involved in sexual behaviour and interindividual or
even interspecies communication. A velum spans the interior of each leg and is formed by cap cells. Three
scolopidia stretch from the velum to the leg wall, each containing one sensory neuron with a dendrite and
attached cilia. The dendrite is accompaigned by a so-called scolopale cell which generates an electron-rich
intracellular structure surrounding the dendrite.[17]
In the cave cricket Troglophilus neglectus the subgenual organ is fairly simple and is associated with an
intermediary organ. Both are innervated either by one or two nerves, depending on the individual animal.[4]
In the splay-footed cricket Comicus calcaris, the subgenual organ is associated with a crista acustica
homolog and an intermediary organ. This organ system is not suitable for hearing sounds, but it is possible
that this organ system formed through reduction of a previously existing hearing organ. All three organs are
innervated by the same nerve and the subgenual organ of this genus has the largest number of nerve cells of
all Ensifera without tympana.[18]
In the heelwalker Karoophasma biedouwense, the subgenual organ is associated with four additional organs
containing scolopidia, a trait shared with Mantophasmatodea. Between 15 and 30 scolopidial cells make up
the subgenual organ, more in the hind limbs. They form a fan-like structure branching out from the anterior
side of the limb. Campaniform sensilla are also associated with the organ. There are no sex differences in the
SG organ. All five organs have less sensitivity for high frequencies and appear to be used for male
specimens at identifying female animals.[19]
In the cricket Gryllus assimilis the sub genual organ fills up most of the tibia and has a fan-like shape. It is
connected to two different neuronal ganglia, one with three bipolar neurons and the other with tens of
neurons that also supplies other insect sensory organs located in the leg. Most of the subgenual organ is
innervated by this major ganglion, except for the more proximal part. There is also an intermediary organ
and a tympanal organ.[20]
In Nezara viridula the organ contains only two scolopidia. The organ is appended to the forward side of the
tibia and hangs into the tibial blood cavity. Two cap cells give it a fan-like shape. Analysis of the neuronal
response to vibration indicates that the organ undergoes resonance after stimulation, only slowly dampening.
The sensitivity is correlated to the insect's own sounds.[21]
In the stick insects Carausius morosus and Siyploidea sipylus, a highly developed distal organ is present in
addition to the subgenual organ, but it contains less scolopidia than the subgenual organ. The organ itself has
a semicircular shape inside the limb and is supplied by three different nerves, one of which also targets the
distal organ. In both species, there are more than 40 scolopidia in the subgenual organ.[22]

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References
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ISSN 0340-7594.
2. Virant-Doberlet, Meta; Cokl, Andrej (2004). "Vibrational communication in insects". Neotropical Entomology. 33
(2): 121134. doi:10.1590/S1519-566X2004000200001. ISSN 1519-566X.
3. R. F. Chapman; Stephen J. Simpson; Angela E. Douglas (January 2013). The Insects: Structure and Function.
Cambridge University Press. p. 752. ISBN 978-0-521-11389-2.
4. Strauss, J.; Stritih, N.; Lakes-Harlan, R. (2014). "The subgenual organ complex in the cave cricket Troglophilus
neglectus (Orthoptera: Rhaphidophoridae): comparative innervation and sensory evolution". Royal Society Open
Science. 1 (2): 140240140240. doi:10.1098/rsos.140240. ISSN 2054-5703.
5. John L. Capinera (11 August 2008). Encyclopedia of Entomology. Springer Science & Business Media. p. 864.
ISBN 978-1-4020-6242-1.
6. Broad, G. R.; Quicke, D. L. J. (2000). "The adaptive significance of host location by vibrational sounding in
parasitoid wasps". Proceedings of the Royal Society B: Biological Sciences. 267 (1460): 24032409.
doi:10.1098/rspb.2000.1298. ISSN 0962-8452.
7. Rssler, Wolfgang (1992). "Postembryonic development of the complex tibial organ in the foreleg of the
bushcricket Ephippiger ephippiger (Orthoptera, Tettigoniidae)". Cell & Tissue Research. 269 (3): 505514.
doi:10.1007/BF00353905. ISSN 0302-766X.
8. Ball, Eldon; Young, David (1974). "Structure and development of the auditory system in the prothoracic leg of
the cricket Teleogryllus commodus (walker)". Zeitschrift fr Zellforschung und mikroskopische Anatomie. 147
(3): 313324. doi:10.1007/BF00307467. ISSN 0302-766X.
9. Kutsch, Wolfram (1989). "Formation of the receptor system in the hind limb of the locust embryo". Roux's
Archives of Developmental Biology. 198 (1): 3947. doi:10.1007/BF00376369. ISSN 0930-035X.
10. J. T. Wong, W. T. Yu & T. P. O'Connor (September 1997). "Transmembrane grasshopper Semaphorin I promotes
axon outgrowth in vivo". Development. 124 (18): 35973607. PMID 9342052.
11. Kilpinen, O.; Storm, J. (1997). "Biophysics of the subgenual organ of the honeybee, Apis mellifera". Journal of
Comparative Physiology A. 181 (4): 309318. doi:10.1007/s003590050117. ISSN 0340-7594.
12. Storm, Jesper; Kilpinen, Ole (1998). "Modelling the subgenual organ of the honeybee, Apis mellifera". Biological
Cybernetics. 78 (3): 175182. doi:10.1007/s004220050424. ISSN 0340-1200.
13. Menzel, Johannes G.; Tautz, Jrgen (1994). "Functional morphology of the subgenual organ of the carpenter ant".
Tissue and Cell. 26 (5): 735746. doi:10.1016/0040-8166(94)90056-6. ISSN 0040-8166.
14. Howse, P.E. (1964). "An investigation into the mode of action of the subgenual organ in the termite,
Zootermopsis angusticollis Emerson, and in the cockroach, Periplaneta americana L.". Journal of Insect
Physiology. 10 (3): 409424. doi:10.1016/0022-1910(64)90065-4. ISSN 0022-1910.
15. Shaw, Stephen R. (1994). "Re-evaluation of the absolute threshold and response mode of the most sensitive
know ?vibration? detector, the cockroach's subgenual organ: A cochlea-like displacement threshold and a direct
response to sound". Journal of Neurobiology. 25 (9): 11671185. doi:10.1002/neu.480250911. ISSN 0022-3034.
16. Moran, David T.; Carter Rowley, J. (1975). "The fine structure of the cockroach subgenual organ". Tissue and
Cell. 7 (1): 91105. doi:10.1016/S0040-8166(75)80009-7. ISSN 0040-8166.
17. Devetak, Duan; Pabst, Maria Anna (1994). "Structure of the subgenual organ in the green lacewing, Chrysoperla
carnea". Tissue and Cell. 26 (2): 249257. doi:10.1016/0040-8166(94)90100-7. ISSN 0040-8166.
18. Strau, Johannes; Lakes-Harlan, Reinhard (2010). "Neuroanatomy of the complex tibial organ in the splay-footed
cricket Comicus calcaris Irish 1986 (Orthoptera: Ensifera: Schizodactylidae)". The Journal of Comparative
Neurology. 518 (22): 45674580. doi:10.1002/cne.22478. ISSN 0021-9967.
19. Eberhard, M.J.B.; Lang, D.; Metscher, B.; Pass, G.; Picker, M.D.; Wolf, H. (2010). "Structure and sensory
physiology of the leg scolopidial organs in Mantophasmatodea and their role in vibrational communication".
Arthropod Structure & Development. 39 (4): 230241. doi:10.1016/j.asd.2010.02.002. ISSN 1467-8039.
20. Friedman, Morton H. (1972). "A light and electron microscopic study of sensory organs and associated structures
in the foreleg tibia of the cricket,Gryllus assimilis". Journal of Morphology. 138 (3): 263327.
doi:10.1002/jmor.1051380302. ISSN 0362-2525.
21. okl, Andrej (1983). "Functional poperties of viboreceptors in the legs ofNezara viridula (L.) (Heteroptera,
Pentatomidae)". Journal of Comparative Physiology A. 150 (2): 261269. doi:10.1007/BF00606376.
ISSN 0340-7594.
22. Strau, Johannes; Lakes-Harlan, Reinhard (2013). "Sensory neuroanatomy of stick insects highlights the
evolutionary diversity of the orthopteroid subgenual organ complex". Journal of Comparative Neurology. 521
(16): 37913803. doi:10.1002/cne.23378. ISSN 0021-9967.

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Categories: Arthropod morphology
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