Alarm
pheromone:
Alarm pheromones
serve to rapidly disperse a group of insects usually as a response to predation
or threat. alarm pheromone, it is generally considered necessary to demonstrate
that (i) the chemical(s) is released exclusively under exposure to hazard (e.g.
predator attack), (ii) the signal is perceived by conspecifics, and (iii) it
induces in the receiving individuals behavioral reactions similar to that
induced when directly exposed to the same danger (Wyatt, 2003). Generally,
adaptive responses to the reception of an alarm pheromone may be classified as
evasive (e.g., receivers flee from the pheromone releaser) or aggressive
(receivers move toward the signal and attack or harass the predator). Observed
reactions can vary according to the concentration of pheromone released and
also with prior experience of the receiver (Howse, 1998).
The alarm pheromones of many arthropods are also used as
defensive allomones, activity inhibitors, cryptic alarm pheromones, aggregative
attractants, robbing agents, digging agents, trail pheromones, and
antimicrobial agents (Blum,1996).
Alarm pheromones have been identified chiefly in three
subfamilies of ants and their distribution appears to be chemosystematically
significant. Myrmicine genera emphasize 3-alkanones as alarm releasers, whereas
methyl ketones, primarily of terpenoidal origin, are widely utilized as alarm
pheromones in the subfamily Dolichoderinae. Formicine species may employ formic
acidas an alarm pheromone in addition to the compounds produced in the
mandibular and Dufour's glands. The mandibular gland pheromones are chiefly
acyclic monoterpene aldehydes (e.g., citronellal) which are relatively low
boiling compounds. Higher boiling n-alkanes are
produced in the Dufour's glands and may serve as more persistent releasers of
alarm behavior. Alarm pheromones as well as the caste-specific pheromones of
male bees and ants, probably also serve as defensive products (Blum and Brand,1972). These kinds of pheromones are usually of short duration and the
dispersed individuals usually reform aggregations. Some individuals release this pheromone to
exhibit aggressive behaviour in the presence of predators. Alarm pheromones have been recorded in the cockroaches,
treehoppers, aphids, bedbugs, termites, and social Hymenoptera ( Kerkut and
Gilbert, 1985; Blum, 1996). The
alram pheromone has also been recorded in fishes (Scholz et al., 2000);
amphibians (Scholz et al.,2000) and in mammals like cows and feathered
minnos (Mathis et al.,1995).
In Homopterans, so far only in two families the alarm pheromones have been
studied. In membracids the body wall
tissue is the source of pheromone (Nault et al., 1974), whereas, in aphids the
cornicles are the source of pheromone (Bowers et al., 1972). In many
other aphids alarm pheromones are reported (Wientjens et al.,1873;Bowers et al.,1972;Edwards
et al.,1973; Phelan et al.,1976; Pickett and Griffiths,1980; Gibson and Pickett,1983). In Hemiptera, four families are identified
possessing alarm pheromone (Blum, 1985).
In Pyrrhocoridae,
the 3rd dorsal abdominal
parts; in Cimicidae the dorsal abdominal gland; in Coreidae the metathoracic
glands in adults and dorsal abdominal gland of nymphs (Blatt et al.,
1998; Leal et al.,1994) have been reported to produce alarm pheromones.
In
Pentatomidae, it is located in adults
metathoracic gland (Ishiwatari,1974). In Acanthosomidae, the alarm pheromone has been reported from
dorsal abdominal gland in the larvae and
metathoracic glands in the adults (Maschwitz and Gutmann, 1979). In Miridae, the metathoracic scent gland is
the source of alarm pheromone (Groot et al .,2001). In the Thysanoptera
(thrips), the alarm pheromone is recorded from anal fluid produced by the
nymphal instars (Teeling et al., 1993).
In Isoptera (termite) the frontal gland of soldiers and homogenial
femuli in workers are found to be the
sources of alarm pheromones (Moore, 1968; Farhat and Iqbal, 1980).
The sources of
alarm pheromone in ants have been located to eleven; the mandibular glands
(Wilson, 1958; Maschwitz, 1964), Pygidal gland (Wilson and Pavan, 1959),
Dufour’s gland and poison gland (Maschwitz, 1974) of metaplural gland of
workers (Masschwitz, 1964), crushed heads of workers (Duffield et al.,
1980). Higher concentration from the heads of aged queens (Francke et al.,
1980), small quantities in the abdomens of aged queen (Hughes et al.,
2001), whole ants with crushed heads, anal gland or some other source of pheromones (Hefetz
and Lloyd, 1983).
In the family
Blattidae of the order Dictioptera, the whole body of the both sexes and glands
associated with the 2nd abdominal spiracle are reported to be the
sources of alarm pheromone (Brossut, 1983; Rollo et al., 1995). Aggressive
behaviour in some of the families of Hymenoptera has also been demonstrated. In Vespidae the poison gland of workers and
females are located to be the source of alarm pheromone (Maschwitz, 1964). Similarly, venom reservoir (Manzoli-Palma et
al., 1998), pupal odour (Landolt et al., 1999) and heads of workers
(Maschwitz, 1984) are identified as the sources of alarm pheromones (Ghent and
Gary, 1962; Maschewitz, 1964; Shearer and Boch, 1965). Even the venom has been identified as the
source of alarm pheromone in honeybees (Veith et al., 1978). Koschewnikow gland is identified as the
sources of alarm pheromones (Lensky et al., 1991). Cassier et al. (1994) reported sting
sheets as a source of alarm pheromone in honeybee. Most Alarm pheromones likely
have evaloved from compound originally having other functions. Specially it has
been proposed that Alarm pheromone may evalve from chemicals used for defense against predators
or from compounds released upon injury (Wyatt, 2003).
The
aggregation pheromone of Wisconsin Ips
pini contains ipsdienol, which is (+)
and (-) enantiomers, and lanierone, and the pheromone of I. grandicollis contains ipsenol.
By contrast, -pinene consistently
enhanced attraction by all three
predators to the pheromones of their Ips prey (Nadir Erbilgin and Kenneth F Raffa, 2001).For a
detailed review of the alarm pheromone, the research work of Blum (1985) may be
consulted.
Similarly 2-heptanone and 2-heptanol are known as ‘alarm’ or repellent
substances in many hymenopterans, including honey bees (Free, 1987; Wongsiri et al., 2006) and several ant species (Vander
Meer et al., 1998). A recently
accepted patent (US patent number 6,071,973) by Vander Meer et al.(2000). In many ants and termites, the glandular sources
of trail and alarm pheromones have been reported (Kaib, 1999; Kaib, 2000;Wyatt,
2003).
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