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''Nasonia vitripennis'' (or ''Mormoniella vitripennis'', or ''Nasonia brevicornis'') is one of four known species under the genus '' Nasonia'' - small parasitoid
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. ...
s that afflict the larvae of parasitic carrion flies such as blowflies and flesh flies, which themselves are parasitic toward nestling birds. It is the best known and most widely studied of the parasitoid wasps, and their study forms a vital part of the information used to describe the order Hymenoptera, along with information from bees and ants. This parasitoid behaviour makes the wasps an interest for the development of
biopesticide A Biopesticide is a biological substance or organism that damages, kills, or repels organisms seens as pests. Biological pest management intervention involves predatory, parasitic, or chemical relationships. They are obtained from organisms inclu ...
and biological systems for controlling unwanted insects. The biosynthetic pathways for sex pheromones in Hymenoptera, determination of sex in development, and many protein and gene product comparisons to other insects have been studied using ''N. vitripennis'' (most often contrasted against the
Western honey bee The western honey bee or European honey bee (''Apis mellifera'') is the most common of the 7–12 species of honey bees worldwide. The genus name ''Apis'' is Latin for "bee", and ''mellifera'' is the Latin for "honey-bearing" or "honey carrying" ...
, ''Apis mellifera''). ''Nasonia vitripennis'' also has a high variety of proteins that have been discovered for venom and detection of odours and has repetitive DNA; this information has been made easier for study since the complete sequencing and release of the genome of N. vitripennis in 2010.


Physiology

As in other ''Nasonia'' wasps, ''N. vitripennis'' is
haplodiploid Haplodiploidy is a sex-determination system in which males develop from unfertilized eggs and are haploid, and females develop from fertilized eggs and are diploid. Haplodiploidy is sometimes called arrhenotoky. Haplodiploidy determines the sex ...
, having haploid males and diploid females, and measures from 2–3 mm in length, with larger and darker-colored females than males. These wasps, like most other insects, show much sexual dimorphism, and females tend to be less easy to distinguish by species than males. ''N. vitripennis'' females have a straight stigmal vein (a short branch from the stigma of the forewing), in comparison to the varying curvature in its three sister species. Males are generally distinguished using antenna and wing shape. Male ''N. vitripennis'' wasps have a spindle-shaped scape (the lower half of the antenna), meaning it is wider in the middle than at either of the joint ends. (This is in comparison to the “angulate” shape seen in ''N. giraulti'' and ''N. oneida'', or the cylindrical shape of ''N. longicornis''). The antennal flagellum is also shorter and wider than in the other three species of ''Nasonia''. Male ''N. vitripennis'' have small forewings, in comparison to other ''Nasonia'' relatives.


Pheromones and sexual behaviour

Male ''N. vitripennis'' wasps produce
pheromones A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
from papillae inside a rectal vesicle, and release pheromones through the anus. Female wasps show no similar organ for pheromone release. Prior research has pointed to the rectal papillae (inside the rectal vesicle) for the purpose of water and electrolyte resorption, since the adult male wasps rarely feed; however, localization techniques, pheromone biosynthesis data and observations of wasp behaviour (tapping abdomen on the ground, leaving traces of pheromone) all point to these organs being used in sexual communication. Cephalic pheromones are also present in ''N. vitripennis'', coming from the mouth of the males during courtship, which females contact with their antennae while signaling their receptivity to mating. The distinctive chemical mixture in male pheromones strongly attracts virgin females; mated females (and females merely exposed to the male pheromones) experimentally have shown little to no attraction. After mating, females then turn their attention to searching for a host larva in which to oviposit – a behaviour which reroutes wasted time that would be spent over-mating to searching for a suitable host. This behavioural ‘switch’ is thought to be caused by pheromones alone, rather than by sperm transfer, as seems to be the case in other insects. Egg-laying in ''N. vitripennis'' occurs in the same fashion in all known species of ''Nasonia'', where a mated female drills into and deposits her eggs under the puparum of fly larvae.


Non-CSD sex determination

Hymenoptera largely reproduce through facultative
parthenogenesis Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and developmen ...
, where haploid males develop from unfertilized eggs (
arrhenotoky Arrhenotoky (from Greek -τόκος ''-tókos'' "birth of -" + ἄρρην ''árrhēn'' "male person"), also known as arrhenotokous parthenogenesis, is a form of parthenogenesis in which unfertilized eggs develop into males. In most cases, parthe ...
) and diploid females develop from fertilized eggs. Sex determination in all ''Nasonia'' has been shown to be due mainly to fertilization status (fertilized or unfertilized as an egg), as well as by chromosome number. ''Nasonia'' and other Chalcid wasps use a different sex determination system than a large portion of Hymenopterans (including
Honey bees A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current cosm ...
), who use CSD (complementary sex determination), where sex is determined by homozygous or heterozygous
alleles An allele (, ; ; modern formation from Greek ἄλλος ''állos'', "other") is a variation of the same sequence of nucleotides at the same place on a long DNA molecule, as described in leading textbooks on genetics and evolution. ::"The chrom ...
of a single gene. The most recent accepted model for this non-CSD system is called Maternal Effect Genomic Imprinting Sex Determination (MEGISD). This model involves a masculinizing/virilizing maternal effect gene that “imprints upon” the cytoplasmic component of oocytes, and an “unimprinted” paternal contribution (in female offspring) that provides a counter effect to
virilization Virilization or masculinization is the biological development of adult male characteristics in young males or females. Most of the changes of virilization are produced by androgens. Virilization is most commonly used in three medical and biology ...
and allows for female development to occur. Since all diploid eggs become female (due to the factor originating in the male genetic contribution that prevents masculinization), this differs from CSD in that under CSD, males can be diploid if they are homozygous or hemizygous.


''Wolbachia''

''
Wolbachia ''Wolbachia'' is a genus of intracellular bacteria that infects mainly arthropod species, including a high proportion of insects, and also some nematodes. It is one of the most common parasitic microbes, and is possibly the most common reproducti ...
'' are maternally inherited, and exist in multiple insect species. Multiple strains of ''Wolbachia'' have been noted in even single strains of ''Nasonia''. These bacteria are a very well-documented
endosymbiont An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within ...
(arguably parasite) of the wasps that can have consequences for fertility when their presence is altered. It has been suggested that ''Wolbachia'' have assisted with genetic divergence through buildup of mutations and selecting for compensatory mechanisms, causing the emergence of new strains to occur more quickly. This is through a model of ‘mitochondrial sweeping’, when ''Wolbachia'' bring mitochondria from one host population to another.


References


External links


Nasoniabase

''Nasonia''
resources hosted at the University of Rochester {{Taxonbar, from=Q13875897 Insects used as insect pest control agents Insects of the United States Pteromalidae