FILE: <bc-10.htm> Pooled References GENERAL INDEX [Navigate to MAIN MENU ]
EXTRANUCLEAR EVENTS INFLUENCING
BEHAVIOR AND
FECUNDITY IN PARASITOIDS AND OTHER
ANIMALS
(Contacts)
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Extranuclear influences on behavior
Direct
Effects of the Cytoplasmic Genome
Chemical Substances Affecting Behavior
Muscidifurax Parasitoid Complex
[ Please
refer also to Selected
Reviews #1, #2
& Detailed Research ]
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Extranuclear influences on behavior involving
cytoplasmic entities are well known among both prokaryotes and eukaryotes
(Beale & Knowles 1978, Cosmides & Tooby 1981, Goodenough 1984, Levine
1973, Sager & Ramanis 1963, Sonneborn 1959), but the subsequent
incorporation of an extranuclear expression into the nuclear genome
apparently have not been found. Extranuclear factors in the form of
microorganisms (e.g., viruses, bacteria, spiroplasmas) can alter sex ratios
in parasitoids by selectively killing developing males or females (Skinner
1982, 1985; Werren et al. 1981, 1986), may confer resistance to host
encapsulation (Krell & Stoltz 1979, Stoltz & Vinson 1977, Stoltz et
al. 1976, Vinson & Stoltz 1986), and affect sex ratios in Drosophila (Poulson &
Sakaguchi 1961), and are passed on to succeeding generations. Cosmides and
Tooby (1981) recently reviewed how cytoplasmic genes control such characters
as allocation of reproductive effort in hermaphrodites, sex ratios of
offspring (Williamson & Poulson 1979), organism size (Faulkner &
Arlett 1964), growth rate, colony size, rate of senescence (Smith &
Rubenstein 1973), competitive ability (Preer et al. 1974), drug resistance in
bacteria, protozoans, fungi, and mammals (Beale & Knowles 1978), and
rates of recombination among nuclear genes (Thoday & Boam 1956). Oishi et
al. (1984) explained how two kinds of microorganisms (spiroplasma and virus)
may interact to modify expressions of the sex-ratio factor in Drosophila. Stoltz & Vinson
(1979) have found viruses in the calyx epithelial cells of endoparasitoids.
Fleming & Summers (1986) found them also in the lumen of the oviduct.
These viruses were passed from parent to offspring, males being able to
transmit viral DNA to females with whom they mated (Stoltz et al. 1986). The direct effects of the cytoplasmic genome on the nuclear
genome has been hypothesized (Cosmides & Tooby 1981), but not
demonstrated. However, the extrachromosomal genetic system can be influenced
by the chromosomal system (Levine 1973). Microorganisms implicated in
inheritance have been known to cause illness and death in male Drosophila (Leventhal 1968). Chemical substances affect behavior of insects following
mating. Reports include a lepidopteran (Webster & Carde 1984), and
ichneumon wasp Venturia (Nemeritis) canescens (Gravenhorst) (where heneicosane was involved)
(Mudd et al. 1982); and the dipteran Drosophila,
involving enzymes (Mane et al. 1983, Richmond & Senior 1981).
Prostaglandins, derivatives of certain polyunsaturated fatty acids, alter egg
laying behavior in crickets (Stanley-Samuelson & Loher 1986). It has been
suggested than an influence on the chemosensory responsiveness of an
individual by chemical cues derived from its parents would be hard to
distinguish from a genetic effect (Corbet 1985). Microorganisms involved in the production of thelytoky
have been identified molecularly by Stouthamer et al. (1993). They comment
that inherited microorganisms are widespread in insects, having been
implicated as causes of female parthenogenesis and cytoplasmic
incompatibility. Normal sexual reproduction can be restored by treatment with
antibiotics. Sequence analysis of the DNA encoding 16S ribosomal RNA shows
that cytoplasmic incompatibility bacteria from diverse insect taxa are
closely related, sharing 95% sequence similarity. They belong to the alpha
subdivision of Proteobacteria. Stouthamer et al. (1993) show that
parthenogenesis-associated bacteria from parasitoid Hymenoptera fall into
this bacterial group, having up to 99% sequence similarity to some
incompatibility microorganisms. Both incompatibility and parthenogenesis
microorganisms alter host chromosome behavior during early mitotic division
in the egg. Incompatibility bacteria act by interfering with paternal
chromosome incorporation in fertilized eggs, while parthenogenesis bacteria
prevent segregation of chromosomes in unfertilized eggs. These traits are
adaptive for the microorganisms. Judging from their sequence similarities,
Stouthamer et al. (1993) concluded that parthenogenesis bacteria and
cytoplasmic incompatibility bacteria form a monophyletic group of
microorganisms that specialize in manipulating chromosome behavior and insect
reproduction. The
Muscidifurax Complex. Research on five distinctive races of the
genus Muscidifurax
(see Research)
has uncovered a polygenic system controlling rates of larval cannibalism and
therefore reproductive success. The genes involved are able to cause partial
expression of the traits they govern shortly after insemination and before
being inherited by the progeny. Extranuclear phases prior to chromosomal
inheritance may involve microorganisms and/or enzymes present in hymenopteran
seminal fluid. The ability to
change expression of a quantitative character immediately after mating,
either positively or negatively, challenges accepted views of polygenic loci,
and it may be that such loci are not in fact inherited, but rather another
group of genes which have the capability to switch on or off the loci. Such
genes may influence DNA methylation of the loci controlling oviposition
behavior, as shown for other organisms (). All polygenic loci may be
perpetually present for a given quantitative trait in all individuals of both
Muscidifurax raptorellus races, but they are
either activated or inactivated by substances under the control of another
group of genes (Legner 1993 ). Allowing natural
selection for nonlethal undesirable and desirable characteristics to begin to
act in the parental generation theoretically accelerates evolution in the Muscidifurax
system. Traits, which are detrimental to the population, might thus be more
prone to elimination and beneficial ones may be expressed in the mother
before the appearance of her active progeny. If a similar genetic system
occurs more generally in Hymenoptera, it might account partially for the
rapid evolution thought to occur in certain groups of Hymenoptera. The
ability of male Hymenoptera to activate heritable behavior in females with
whom they mate has practical significance in biological control. Greater
importance may be placed on liberated males during mass release strategies
that seek to accelerate and increase the magnitude of parasitism, because it
is possible to convey certain desirable strain characteristics directly to
unmated females already resident in the environment. This subject will be
treated in greater detail in a subsequent section on arrhenotoky. Studies with new field isolates
of a Peruvian strain in 1995 by Richard Stouthamer et al. (unpublished) have
shown a greater involvement of larval cannibalism and much complexity in this
species' reproduction. Indeed
survival mechanisms in parasitoids include many behaviors; among which
increased cannibalism by more aggressive larvae may be triggered during times
of host scarcity. An account may be found in <aggress.htm>. [Please refer
also to Related Research ] Exercise 10.1. How may you distinguish behavior that is
regulated by extranuclear factors from that regulated by genetic factors? Exercise 10.2. Name the kinds of organisms that have been
implicated in triggering extranuclear responses in animals. Exercise 10.3. Could extranuclear factors be used to
control behavior in insects? REFERENCES: [Additional references may be
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3: 175-208. |