Biology Reference
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seems that the tick and the symbiont have not coevolved, but the bacteria are
vertically transmitted from females to their progeny and are transmitted trans-
stadially. Pistone et al. (2012) suggest, “The high prevalence of M. mitochondrii
in wild-collected adult females of I. ricinus seems to suggest a beneficial role .
However, the decrease in the number of bacteria in laboratory-reared ticks,
without visible effects on the biology of the hosts, suggests the presence of the
symbionts, if beneficial, is facultative, at least in laboratory conditions.”
Antlions (Myrmeleontidae) suck out the body fluid after first paralyzing their
prey with a toxin produced by salivary gland secretions produced by bacteria
located in the salivary glands. The paralyzing toxin produced by these bacterial
endosymbionts is a homolog of GroEL, a heat-shock protein that functions as
a molecular chaperone in E. coli ( Yoshida et al. 2001 ). In the antlion, the GroEL
protein may act on receptors in prey insects to induce paralysis. The antlion
symbionts perhaps evolved this nonchaperone function to establish a mutually
beneficial antlion-bacterium relationship. Yoshida et  al. (2001) speculated that
insecticidal proteins may be produced by other endosymbionts to help addi-
tional fluid-feeding predatory insects.
4.12.2 Wolbachia and Cardinium
Wolbachia , a genus of α -proteobacteria, commonly are found in arthropods
( Werren et  al. 1995, Rigaud and Rousset 1996, O'Neill et  al. 1997, Jeyaprakash
and Hoy 2000 ). Wolbachia are intracellular gram-negative rods that cannot be
cultured easily outside their hosts. Wolbachia infection rates range from 17 to
76% of all arthropod species ( Werren et  all 1995, Jeyaprakash and Hoy 2000 ).
Wolbachia also have been found in crustaceans ( Rigaud 1999 ) and nematodes
( Bandi et al. 1999, Bazzocchi et al. 2000 ). An understanding of their physiologi-
cal and phenotypic effects on their hosts is still being developed. The phylogeny
of Wolbachia in nematodes is congruent with the phylogeny of their hosts, sug-
gesting they share a long coevolutionary history. Treatment with the antibiotic
tetracycline inhibits normal reproduction and development of filarial nematodes
that harbor Wolbachia , suggesting that Wolbachia are necessary to the nema-
tode ( Langworthy et al. 2000 ). That is not true for arthropods, where it seems a
great deal of horizontal transfer has occurred.
Wolbachia have been implicated as both the cause of alterations in sex ratio
(resulting in thelytoky and male killing), to be discussed further in the Chapter
10 on sex determination, and cytoplasmic incompatibility in arthropods. Some
Wolbachia improve fertility or vigor, whereas others seem to decrease these
traits in their hosts. The molecular mechanism(s) by which reproductive incom-
patibility are induced by Wolbachia are hypothesized to be due to Wolbachia 's
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