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The osmeterium is a defensive organ found in all papilionid larvae, in all stages. 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, small spiders and mantids. 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 In chemistry, an ester is a chemical compound, compound derived from an acid (either organic or inorganic) in which the hydrogen atom (H) of at least one acidic hydroxyl group () of that acid is replaced by an organyl group (R). These compounds c ...
. 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, and storage of the osmeterial secretion.


Structure

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. 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.


Osmeterial secretion

The osmetrial secretion consists of a mixture of volatile organic acids, such as
isobutyric acid Isobutyric acid, also known as 2-methylpropanoic acid or isobutanoic acid, is a carboxylic acid with structural formula ( CH3)2CH COOH. It is an isomer of butyric acid. It is classified as a short-chain fatty acid. Deprotonation or esterificati ...
and
2-methylbutyric acid 2-Methylbutanoic acid, also known as 2-methylbutyric acid is a branched-chain alkyl carboxylic acid with the chemical formula CH3CH2CH(CH3)CO2H, classified as a short-chain fatty acid. It exists in two enantiomeric forms, (''R'')- and (''S'')-2-m ...
. The secretion of '' Papilio demodocus'' pre-final
larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...
l
instar An instar (, from the Latin '' īnstar'' 'form, likeness') is a developmental stage of arthropods, such as insects, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...
contains methyl 3-hydroxybutanoate, 3-hydroxybutanoic acid, α-pinene, myrcene,
limonene Limonene () is a colorless liquid aliphatic hydrocarbon classified as a cyclic monoterpene, and is the major component in the essential oil of citrus fruit peels. The (+)-isomer, occurring more commonly in nature as the fragrance of oranges, ...
, β-phellandrene, (''Z'')-ocimene, (''E'')-ocimene, β-caryophyllene, (''E'')-β-farnesene, and germacrene A, as well as a number of unidentified sesquiterpenoids. and that of its final instar contains 2-methylpropanoic acid, 2-methylbutanoic acid, and their methyl and ethyl esters as major constituents. Volatile components of the secretion of '' Parnassius glacialis'' ( Parnassiinae, Parnassiini) consisted of
isobutyric acid Isobutyric acid, also known as 2-methylpropanoic acid or isobutanoic acid, is a carboxylic acid with structural formula ( CH3)2CH COOH. It is an isomer of butyric acid. It is classified as a short-chain fatty acid. Deprotonation or esterificati ...
,
2-methylbutyric acid 2-Methylbutanoic acid, also known as 2-methylbutyric acid is a branched-chain alkyl carboxylic acid with the chemical formula CH3CH2CH(CH3)CO2H, classified as a short-chain fatty acid. It exists in two enantiomeric forms, (''R'')- and (''S'')-2-m ...
, and their methyl esters. That of '' Sericinus montela'' ( Parnassiinae, Zerynthiini) was characterized as
monoterpene Monoterpenes are a class of terpenes that consist of two isoprene units and have the molecular formula C10H16. Monoterpenes may be linear (acyclic) or contain rings (monocyclic and bicyclic). Modified terpenes, such as those containing oxygen func ...
hydrocarbon In organic chemistry, a hydrocarbon is an organic compound consisting entirely of hydrogen and carbon. Hydrocarbons are examples of group 14 hydrides. Hydrocarbons are generally colourless and Hydrophobe, hydrophobic; their odor is usually fain ...
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. 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. *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'', '' P. demodocus'' 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

Eversion of the osmeterial gland is demonstrably effective in reducing predation by ants and small spiders, and its chemical constituents have been shown to repel or kill ants,HONDA, KEIICHI. Defensive potential of components of the larval osmeterial secretion of papilionid butterflies against ants. Physiological Entomology - Blackwell Publishing Ltd

DO - 10.1111/j.1365-3032.1983.tb00346.x
and mantids.


References


External links

{{commons category
Tree of Life
Papilionidae Insect anatomy