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{{short description|Odoriferous defensive structure}}
[[Image:Osmeterium1.JPG|thumb|right|Close up of the osmeterium]]
[[File:Papilio xuthus Larva 2011-07-01.jpg|thumb|{{center|Osmeterium of ''[[Papilio xuthus]]'' Larva. <br>Upper: Osmeterium partly everted. Lower: undisturbed}}]]
The osmeterium is a defensive organ found in all [[Papilionidae|Papilionid]] larva, in all stages<ref name=Chattopadhyay>Chattopadhyay, J. 2011. The structure and defensive efficacy of glandular secretion of the larval osmeterium in Graphium agamemnon agamemnon Linnaeus, 1758 (Lepidoptera: Papilionidae)</ref>. The organ is situated in the [[prothorax|prothoracic]] segment and can be everted when the larva feels threatened. When not everted the organ is stored inverted within the body. The everted organ resembles a fleshy forked tongue not unlike a snake tongue and this along with the large eye like spots on the body might be used to startle birds and small reptiles. The osmeterial organ also exudes a foul smelling liquid which serves to repel [[ants]], small [[spiders]] and [[mantis|mantids]].
The '''osmeterium''' is a defensive organ found in all [[Papilionidae|papilionid]] larvae, in all stages.<ref name=Chattopadhyay/> The organ is situated in the [[prothorax|prothoracic]] segment and can be everted when the larva feels threatened. The everted organ resembles a fleshy forked tongue (not unlike a snake tongue), and this along with the large eye-like spots on the body might be used to startle birds and small reptiles. The osmeterial organ remains inside the body in the thoracic region in an inverted position and is everted when the larva is disturbed in any way emitting a foul, disagreeable odor which serves to repel [[ants]],<ref name=eisner_meinwald_1965/> small [[spiders]]<ref name=Damman_1986/> and [[mantis|mantids]].<ref name=Chow_Tsai_1989/> To humans, this odour is rather strong but not unpleasant, usually smelling like a concentrated scent of the caterpillar's food plant and pineapple.


The constitution of the osmeterial secretion varies from species to species but on the whole is made up of one of the following
The constitution of the [[#osmetrial secretion|osmeterial secretion]] varies from species to species and contains [[monoterpene|monoterpene hydrocarbons]], [[sesquiterpene|sesquiterpenic compounds]] or a mixture of [[Aliphatic compound#Aliphatic acids|aliphatic acids]] and [[esters]].
*[[monoterpene|monoterpene hydrocarbons]]
*[[sesquiterpene|sesquiterpenic compounds]]
*a mixture of [[Aliphatic_compound#Aliphatic_acids|aliphatic acids]] and [[esters]]
It has been found that in certain species the constituents of the secretion varies between the 4th instar and the 5th instar. Based on this the papilionid species can be grouped into 2 sets. One where the quality of the secretion varies and the other where it does not.<ref name=honda_secretion>HONDA K. 1980: Osmeterial secretions of papilionid larvae in the genera Luehdorfia, Graphium and Atrophaneura (Lepidoptera). Insect Biochem. 10: 583-588.</ref>


The fine structure of the osmetrium of <i>Papilio demoleus libanius</i> Fruhstorfer has been studied and found to contain 2 types of cells
The fine structure of the osmeterium of ''[[Papilio demoleus]] libanius'' Fruhstorfer has been studied and found to contain 3 types of specialised cells for synthesis, acid secretion,<ref name=ultrastructure/> and storage<ref name=finestructure/> of the osmeterial secretion.
#Ellipsoid gland cells which might be organic acid-secreting cells
⚫
#Tubular arm cells which might be tasked with synthesis and secretion of the mature secretion<ref name=ultrastructure>Crossley, A.C. and Waterhouse, D.F. 1969. The infrastructure of osmeterium and the nature of its secretion in Papilio larvae (Lepidoptera; Papilionidae). Tissue & Cell 1: 525-554.</ref>
⚫
#A third, Slack cells which might serve as storage cells, has also been reported. <ref name=finestructure>Lu, Chow-Chin; Yien Shing Chow (1991) Fine structure of the larval osmeterium of ''Papilio demoleus libanius'' (Lepidoptera: papilionidae). Ann. Entomol. Soc. Am. 84(3):294-302.</ref>


==Structure==
==Structure==
[[File:Papilio demodocus larva IMG 8543s.jpg|thumb|''[[Papilio demodocus]]'' larva everting its osmeterium fully, and with one horn applying the repugnatorial secretion to the finger that irritated it]]
⚫
The fine structure of the osmeterium gland of Papilio larvae consists of 2 main cell types.
⚫
*Ellipsoid gland cells have an extensively infolded basal plasma membrane, abundant ribosomes and whorls of smooth endoplasmic reticulum. The apical plasma membrane bears long microvilli extending into a mass of granular material containing electron-lucid cavities. Tangential slits occur in the epicuticle. These cells are presumed to be organic acid-secreting cells.
⚫
*Tubular arm cells contain heterogeneous, electron-dense inclusions, extensively-branched nuclei and large mitochondria sometimes distended with electron-dense material. The apical plasma membrane bears short microvilli. The inner, dense epicuticle forms a complex ramifying system. These glandular cells presumably synthesize and secrete the mature secretion.<ref name=ultrastructure/>
⚫
Specific study of [[Papilio demoleus|Papilio demoleus libanius]] Fruhstorfer shows the possible existence of a third type of cell
⚫
*Slack cells, reported for the first time, loosely surround the ellipsoid gland cells. These cells possess papillae, well-developed, rough endoplasmic reticulum, and numerous secretory vacuoles of various sizes and electron density. They may function as storage cells. <ref name=finestructure/>


⚫
The fine structure of the osmeterium gland of ''Papilio'' larvae consists of the following cell types.
⚫
==Osmetrial secretion==
⚫
*Ellipsoid gland cells have an extensively infolded basal [[plasma membrane]], abundant [[ribosome]]s and whorls of [[smooth endoplasmic reticulum]]. The [[Apical (anatomy)|apical]] plasma membrane bears long [[microvilli]] extending into a mass of granular material containing electron-lucid cavities. Tangential slits occur in the [[epicuticle]]. These cells are presumed to be organic acid-secreting cells.
⚫
Studies on osmeterial secretion makes it possible to classify the Papilionid species into at least two large categories in terms of the chemical property of osmeterial secretion.
⚫
*Tubular arm cells contain heterogeneous, electron-dense inclusions, extensively-branched nuclei and large [[mitochondria]], sometimes distended with electron-dense material. The apical plasma membrane bears short microvilli. The inner, dense epicuticle forms a complex ramifying system. These glandular cells presumably synthesize and secrete the mature secretion.<ref name=ultrastructure/>
⚫
*One is a group in which the chemical constitution of osmeterial secretion of the last larval instar markedly differs in quality from those of the younger larvae. The results derived from [[Papilio protenor]] (HONDA, 1980), [[Papilio demodocus|P. demodocus]] (BURGER et al., 1978) and other Papilio species ([[Papilio helenus|P. helenus]], [[Papilio machaon|P. machaon]], [[Papilio memnon|P. memnon]], [[Papilio bianor|P. bianor]], [[Papilio maackii|P. maccki]], [[Papilio xuthus|P. xuthus]], etc., unpublished work) may assign the genus Papilio (tribe Papilionini) to this group, which can be termed 'heterogeneous type'.
⚫
Specific study of [[Papilio demoleus|''Papilio demoleus libanius'']] Fruhstorfer shows the possible existence of a third type of cell
⚫
*The genera [[Luehdorfia]] (tribe [[Zerynthiini]]), [[Graphium (butterfly)|Graphium]] (tribe [[Graphiini]]) and [[Atrophaneura]] (tribe [[Troidini]]) apparently belong to the other group in which no qualitative change of osmeterial secretion occurs at the last larval ecdysis. This group can be designated as 'homogeneous type', which is further subdivisible into three types. <ref name=honda_secretion/><ref>Eisner, T., and Y.C. Meinwald. 1965. The defensive secretions of a caterpillar (Papilio). Science 150: 1733-1735.</ref>
⚫
*Slack cells, loosely surround the ellipsoid gland cells. These cells possess papillae, well-developed, rough endoplasmic reticulum, and numerous secretory vacuoles of various sizes and electron density. They may function as storage cells.<ref name=finestructure/>

⚫
==Osmeterial secretion==
The osmetrial secretion consists of a mixture of volatile [[organic acids]], such as [[isobutyric acid]] and [[2-methylbutyric acid]].<ref name=Chow_Tsai_1989/>

The secretion of ''[[Papilio demodocus]]'' pre-final [[larva]]l [[instar]] contains methyl 3-hydroxybutanoate, [[3-hydroxybutanoic acid]], [[α-pinene]], [[myrcene]], [[limonene]], [[β-phellandrene]], [[ocimene|(''Z'')-ocimene]], (''E'')-ocimene, [[β-caryophyllene]], [[farnesene|(''E'')-β-farnesene]], and [[germacrene A]], as well as a number of unidentified [[sesquiterpenoid]]s.<ref name="Burger_P_demodocus_1985"/> and that of its final instar contains [[2-methylpropanoic acid]], 2-methylbutanoic acid, and their methyl and ethyl esters as major constituents.<ref name="Burger_P_demodocus_1978"/>

Volatile components of the secretion of ''[[Parnassius glacialis]]'' ([[Parnassiinae]], [[Parnassiini]]) consisted of [[isobutyric acid]], [[2-methylbutyric acid]], and their methyl esters. That of ''[[Sericinus montela]]'' ([[Parnassiinae]], [[Zerynthiini]]) was characterized as [[monoterpene]] [[hydrocarbon]]s comprising β-myrcene (major), α-pinene, [[sabinene]], limonene, and β-phellandrene, and of ''[[Pachliopta aristolochiae]]'' ([[Papilioninae]], [[Troidini]]) was composed of numerous sesquiterpene hydrocarbons, including [[α-himachalene]], α-amorphene, and germacrene A, and a few oxygenated sesquiterpenoids.<ref name=honda_1995/>

⚫
Studies on osmeterial secretion makes it possible to classify the Papilionid species into two large categories on the basis of the chemical property of osmeterial secretion.<ref name=honda_1980/>
⚫
*One is a group in which the chemical constitution of osmeterial secretion of the last larval instar markedly differs in quality from those of the younger larvae. The results derived from ''[[Papilio protenor]]'',<ref name=honda_1980/> ''[[Papilio demodocus|P. demodocus]]'' <ref name="Burger_P_demodocus_1978"/> and other ''Papilio'' species (''[[Papilio helenus|P. helenus]]'', ''[[Papilio machaon|P. machaon]]'', ''[[Papilio memnon|P. memnon]]'', ''[[Papilio bianor|P. bianor]]'', ''[[Papilio maackii|P. maccki]]'', ''[[Papilio xuthus|P. xuthus]]'', etc., unpublished work) may assign the genus ''Papilio'' (tribe Papilionini) to this group, which can be termed 'heterogeneous type'.
⚫
*The genera ''[[Luehdorfia]]'' (tribe [[Zerynthiini]]), ''[[Graphium (butterfly)|Graphium]]'' (tribe [[Graphiini]]) and ''[[Atrophaneura]]'' (tribe [[Troidini]]) apparently belong to the other group, in which no qualitative change of osmeterial secretion occurs at the last larval ecdysis. This group can be designated as 'homogeneous type', which is further subdivisible into three types.
**Monoterpene - ''Luehdorfia'' (Zerynthiini, Parnassiinae)
**Sesquiterpene - ''Atrophaneura'' (Troidini, Papilionnae)
**Aliphatic acid and ester - ''Graphium'' (Graphiini, Papilioninae)


==Function==
==Function==
Eversion of the osmeterial gland is demonstrably effective in reducing predation by ants and small spiders (Damman, 1986), and its chemical constituents have been shown to repel or kill ants (Eisner and Meinwald, 1965; Honda, 1983) and mantids (Chow and Tsai, 1989)<ref name=Chattopadhyay/>
Eversion of the osmeterial gland is demonstrably effective in reducing predation by ants and small spiders,<ref name=Damman_1986/> and its chemical constituents have been shown to repel or kill ants,<ref name=eisner_meinwald_1965/><ref name="Honda 1983">HONDA, KEIICHI. Defensive potential of components of the larval osmeterial secretion of papilionid butterflies against ants. Physiological Entomology - Blackwell Publishing Ltd. [https://dx.doi.org/10.1111/j.1365-3032.1983.tb00346.x] DO - 10.1111/j.1365-3032.1983.tb00346.x</ref> and mantids.<ref name=Chattopadhyay/><ref name=Chow_Tsai_1989/>


==References==
==References==
{{reflist|refs=
<references/>
⚫
<ref name=finestructure>Lu, Chow-Chin; Yien Shing Chow (1991) Fine structure of the larval osmeterium of ''Papilio demoleus libanius'' (Lepidoptera: papilionidae). Ann. Entomol. Soc. Am. 84(3):294-302.</ref>
⚫
<ref name=ultrastructure>Crossley, A.C. and Waterhouse, D.F. 1969. The ultrastructure of the osmeterium and the nature of its secretion in Papilio larvae (lepidoptera). Tissue & Cell 1: 525-554.</ref>
<ref name=Chattopadhyay>Chattopadhyay, J. 2011. The structure and defensive efficacy of glandular secretion of the larval osmeterium in Graphium agamemnon agamemnon Linnaeus, 1758 (Lepidoptera: Papilionidae).</ref>
<ref name=honda_1980>HONDA K. 1980: Osmeterial secretions of papilionid larvae in the genera Luehdorfia, Graphium and Atrophaneura (Lepidoptera). Insect Biochem. 10: 583-588.</ref>
<ref name="Burger_P_demodocus_1978">Burger, B.V., Roth, M., Le Roux, M., Spies, H.S.C., Truter, V. and Geertsema, H. 1978. The chemical nature of the defensive larval secretion of the citrus Swallowtail, Papilio demodocus. J. Insect Physiol. 24: 803-805.</ref>
<ref name="Burger_P_demodocus_1985">Burger, B.V., Munro, Z., Roth, M., Spies, H.S.C., Truter, V. and Greetsema, H. 1985. Constituents of osmeterial secretion of prefinal instar larvae of citrus swallowtail, Papilio demodocus (Esper) (Lepidoptera: Papilionidae). J. Chem. Ecol. 11 (8): 1093-1114.</ref><ref name=Chow_Tsai_1989>Chow, Y.S. and Tsai, R.S. 1989. Protective chemicals in caterpillar survival. Experientia (Basel) 45 (4): 390-392.</ref>
<ref name=Damman_1986>Damman, H. 1986. The osmeterial glands of the swallowtail butterfly Eurytides Marcellus as a defense against natural enemies. Ecol.Entomol. 11: 261-265.</ref>
<ref name=eisner_meinwald_1965>Eisner, T., and Meinwald, Y.C. 1965. The defensive secretions of a caterpillar (Papilio). Science, N.Y. 150: 1733-1735.</ref>
<ref name=honda_1995>Keiichi Honda, Nanao Hayashi. Chemical nature of larval osmeterial secretions of papilionid butterflies in the genera Parnassius, Sericinus and Pachliopta. Journal of Chemical Ecology June 1995, Volume 21, Issue 6, pp 859-867.</ref>
}}


==External links==
==External links==

Latest revision as of 15:51, 22 December 2023

Osmeterium of Papilio xuthus Larva.
Upper: Osmeterium partly everted. Lower: undisturbed

The osmeterium is a defensive organ found in all papilionid larvae, in all stages.[1] The organ is situated in the prothoracic segment and can be everted when the larva feels threatened. The everted organ resembles a fleshy forked tongue (not unlike a snake tongue), and this along with the large eye-like spots on the body might be used to startle birds and small reptiles. The osmeterial organ remains inside the body in the thoracic region in an inverted position and is everted when the larva is disturbed in any way emitting a foul, disagreeable odor which serves to repel ants,[2] small spiders[3] and mantids.[4] To humans, this odour is rather strong but not unpleasant, usually smelling like a concentrated scent of the caterpillar's food plant and pineapple.

The constitution of the osmeterial secretion varies from species to species and contains monoterpene hydrocarbons, sesquiterpenic compounds or a mixture of aliphatic acids and esters.

The fine structure of the osmeterium of Papilio demoleus libanius Fruhstorfer has been studied and found to contain 3 types of specialised cells for synthesis, acid secretion,[5] and storage[6] of the osmeterial secretion.

Structure

[edit]
Papilio demodocus larva everting its osmeterium fully, and with one horn applying the repugnatorial secretion to the finger that irritated it

The fine structure of the osmeterium gland of Papilio larvae consists of the following cell types.

  • Ellipsoid gland cells have an extensively infolded basal plasma membrane, abundant ribosomes and whorls of smooth endoplasmic reticulum. The apical plasma membrane bears long microvilli extending into a mass of granular material containing electron-lucid cavities. Tangential slits occur in the epicuticle. These cells are presumed to be organic acid-secreting cells.
  • Tubular arm cells contain heterogeneous, electron-dense inclusions, extensively-branched nuclei and large mitochondria, sometimes distended with electron-dense material. The apical plasma membrane bears short microvilli. The inner, dense epicuticle forms a complex ramifying system. These glandular cells presumably synthesize and secrete the mature secretion.[5]

Specific study of Papilio demoleus libanius Fruhstorfer shows the possible existence of a third type of cell

  • Slack cells, loosely surround the ellipsoid gland cells. These cells possess papillae, well-developed, rough endoplasmic reticulum, and numerous secretory vacuoles of various sizes and electron density. They may function as storage cells.[6]

Osmeterial secretion

[edit]

The osmetrial secretion consists of a mixture of volatile organic acids, such as isobutyric acid and 2-methylbutyric acid.[4]

The secretion of Papilio demodocus pre-final larval instar contains methyl 3-hydroxybutanoate, 3-hydroxybutanoic acid, α-pinene, myrcene, limonene, β-phellandrene, (Z)-ocimene, (E)-ocimene, β-caryophyllene, (E)-β-farnesene, and germacrene A, as well as a number of unidentified sesquiterpenoids.[7] and that of its final instar contains 2-methylpropanoic acid, 2-methylbutanoic acid, and their methyl and ethyl esters as major constituents.[8]

Volatile components of the secretion of Parnassius glacialis (Parnassiinae, Parnassiini) consisted of isobutyric acid, 2-methylbutyric acid, and their methyl esters. That of Sericinus montela (Parnassiinae, Zerynthiini) was characterized as monoterpene hydrocarbons comprising β-myrcene (major), α-pinene, sabinene, limonene, and β-phellandrene, and of Pachliopta aristolochiae (Papilioninae, Troidini) was composed of numerous sesquiterpene hydrocarbons, including α-himachalene, α-amorphene, and germacrene A, and a few oxygenated sesquiterpenoids.[9]

Studies on osmeterial secretion makes it possible to classify the Papilionid species into two large categories on the basis of the chemical property of osmeterial secretion.[10]

  • One is a group in which the chemical constitution of osmeterial secretion of the last larval instar markedly differs in quality from those of the younger larvae. The results derived from Papilio protenor,[10] P. demodocus [8] and other Papilio species (P. helenus, P. machaon, P. memnon, P. bianor, P. maccki, P. xuthus, etc., unpublished work) may assign the genus Papilio (tribe Papilionini) to this group, which can be termed 'heterogeneous type'.
  • The genera Luehdorfia (tribe Zerynthiini), Graphium (tribe Graphiini) and Atrophaneura (tribe Troidini) apparently belong to the other group, in which no qualitative change of osmeterial secretion occurs at the last larval ecdysis. This group can be designated as 'homogeneous type', which is further subdivisible into three types.
    • Monoterpene - Luehdorfia (Zerynthiini, Parnassiinae)
    • Sesquiterpene - Atrophaneura (Troidini, Papilionnae)
    • Aliphatic acid and ester - Graphium (Graphiini, Papilioninae)

Function

[edit]

Eversion of the osmeterial gland is demonstrably effective in reducing predation by ants and small spiders,[3] and its chemical constituents have been shown to repel or kill ants,[2][11] and mantids.[1][4]

References

[edit]
  1. ^ a b Chattopadhyay, J. 2011. The structure and defensive efficacy of glandular secretion of the larval osmeterium in Graphium agamemnon agamemnon Linnaeus, 1758 (Lepidoptera: Papilionidae).
  2. ^ a b Eisner, T., and Meinwald, Y.C. 1965. The defensive secretions of a caterpillar (Papilio). Science, N.Y. 150: 1733-1735.
  3. ^ a b Damman, H. 1986. The osmeterial glands of the swallowtail butterfly Eurytides Marcellus as a defense against natural enemies. Ecol.Entomol. 11: 261-265.
  4. ^ a b c Chow, Y.S. and Tsai, R.S. 1989. Protective chemicals in caterpillar survival. Experientia (Basel) 45 (4): 390-392.
  5. ^ a b Crossley, A.C. and Waterhouse, D.F. 1969. The ultrastructure of the osmeterium and the nature of its secretion in Papilio larvae (lepidoptera). Tissue & Cell 1: 525-554.
  6. ^ a b Lu, Chow-Chin; Yien Shing Chow (1991) Fine structure of the larval osmeterium of Papilio demoleus libanius (Lepidoptera: papilionidae). Ann. Entomol. Soc. Am. 84(3):294-302.
  7. ^ Burger, B.V., Munro, Z., Roth, M., Spies, H.S.C., Truter, V. and Greetsema, H. 1985. Constituents of osmeterial secretion of prefinal instar larvae of citrus swallowtail, Papilio demodocus (Esper) (Lepidoptera: Papilionidae). J. Chem. Ecol. 11 (8): 1093-1114.
  8. ^ a b Burger, B.V., Roth, M., Le Roux, M., Spies, H.S.C., Truter, V. and Geertsema, H. 1978. The chemical nature of the defensive larval secretion of the citrus Swallowtail, Papilio demodocus. J. Insect Physiol. 24: 803-805.
  9. ^ Keiichi Honda, Nanao Hayashi. Chemical nature of larval osmeterial secretions of papilionid butterflies in the genera Parnassius, Sericinus and Pachliopta. Journal of Chemical Ecology June 1995, Volume 21, Issue 6, pp 859-867.
  10. ^ a b HONDA K. 1980: Osmeterial secretions of papilionid larvae in the genera Luehdorfia, Graphium and Atrophaneura (Lepidoptera). Insect Biochem. 10: 583-588.
  11. ^ HONDA, KEIICHI. Defensive potential of components of the larval osmeterial secretion of papilionid butterflies against ants. Physiological Entomology - Blackwell Publishing Ltd. [1] DO - 10.1111/j.1365-3032.1983.tb00346.x
[edit]
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