Trichogramma

Trichogramma
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Suborder: Apocrita
Superfamily: Chalcidoidea
Family: Trichogrammatidae
Genus: Trichogramma
Species

230+, see text

Trichogramma are minute polyphagous wasps that are endoparasitoids of insect eggs.[1] Trichogramma is one of around 80 genera from the family Trichogrammatidae, with over 200 species worldwide.[2][3][4]

Although there are several groups of egg parasitoids commonly employed for biological control throughout the world, Trichogramma have been the most extensively studied.[5] There have been more than a thousand papers published on Trichogramma and they are the most used biological control agents in the world.[6] Trichogramma are unique to the size limit of how small an insect can be, which would be determined by how few neurons they can fit in their central nervous system, yet exhibit a complex behavior to sustain their life. Trichogramma have less than 10,000 neurons, which is a hundred times fewer than the next smallest insect.[7]

Parasitism

To locate host eggs, adult females use chemical and visual signals, such as egg shape and colour.[6] After she finds a suitable egg, an experienced female will attempt to determine if the egg has previously been parasitized using her ovipositor and antennal drumming (tapping on the egg surface). Females also use antennal drumming to determine the size and quality of the target egg, which determines the number of eggs the female will insert.[8] A single female can parasitize one to ten host eggs a day.

Identification

Trichogramma are small and very uniform in structure which causes difficulty in identifying the separate species.[9][10] As females are all relatively similar, taxonomists rely upon examination of males to tell the different species apart using features of their antennae and genitalia.[11][12]

The first description of a Trichogramma species was in North America in 1871 by Charles V. Riley. He described the tiny wasps that emerged from eggs of the Viceroy butterfly as Trichogramma minutum.[3] In taxonomy, original specimens are very important as they are the basis of reference for subsequent descriptions of species. The original specimens, however, were lost. Riley also described a second species in 1879 as Trichogramma pretiosum, but these specimens were also lost. To correct these errors, entomologists returned to the areas where Riley originally found the species and obtained neotype specimens of T. minutum and T. pretiosum. These specimens are now preserved properly in the United States National Museum.[3] Currently the number of Trichogramma species is over 200 but as of 1960 only some 40 species of Trichogramma had been described.[13]

Wolbachia in Trichogramma

Wolbachia is a widespread bacterium that infects insect's organs, most commonly, the reproductive organs.[14] Wolbachia has been observed to alter the host’s reproductive success upon infection.[14] Through a series of manipulations, Wolbachia-infected hosts transmit this intracellular bacterium to uninfected individuals.[14][15] These manipulations include male killing (increasing ratio of infected females who can reproduce), feminization (males become fertile females), parthenogenesis, and cytoplasmic incompatibility.[15] Horizontal transfer of parthenogenesis-inducing Wolbachia, which has been observed in Trichogramma wasps, causes infected females to asexually produce fertile females and nonfunctional males .[16] The effects of this include potential speciation of Trichogramma if Wolbachia is maintained long enough for genetic divergence to occur and for a new species of asexual wasps to become reproductively isolated.[16]

Transmission of the bacterium through horizontal transfer has been observed within the same species and among different species of Trichogramma, including T. kaykai, T. deion, T. pretiosum, T. atopovirilia; however, there are limitations to transmission.[15] In-vitro successful horizontal transfer is uncommon within Trichogramma, which suggests that the density of Wolbachia must be relatively high inside of the host’s ovaries.[15] Cytoplasmic incompatibility of the host and bacterium can also be the source of this unsuccessful transfer in-vitro.[15] These limitations in-vitro suggest that in nature, horizontal transfer by parthenogenesis-inducing Wolbachia may be a difficult and rare phenomenon. However, when looking at the Wolbachia-host associations, the Trichogramma-Wolbachia form a monophyletic group based on several Wolbachia-specific genes, which may be explained by horizontal transfer of Wolbachia between different species.[15] Therefore, although interspecific horizontal transfer of Wolbachia is limited in-vitro, it is likely to occur quite frequently in nature and is not well understood yet.

The effects of Wolbachia in Trichogramma have several evolutionary implications. Commonly, uninfected wasps are unable to breed with infected wasps.[17] Many generations of reproductive isolation of these different groups may result in speciation.[17] In addition, some hosts can evolve with a dependency on Wolbachia for core reproductive functions such as oogenesis, so that eventually an infection is a requirement for successful reproduction.[17] Finally, Wolbachia can influence gender determination in its hosts so that more females are successfully born. This results in a reversal in sexual selection where females must compete for male mates, which has evolutionary implications as it exposes different phenotypes to natural selection.[17]

Biological control

Trichogramma have been used for control of lepidopteran pests for many years. They can be considered the Drosophila of the parasitoid world as they have been used for inundative releases and much of our understanding today comes from experiments with these wasps.[18]

Entomologists in the early 1900s began to rear Trichogramma for biological control. Trichogramma minutum is one of the most commonly found species in Europe and was first mass reared in 1926 on eggs of Sitotroga cerealella.[19]

Nine species of Trichogramma are produced commercially in insectaries around the world with 30 countries releasing them. Trichogramma are used for control on numerous crops and plants, these include cotton, sugarcane, vegetables, sugar beets, orchards and forests.[20] Some of the pests that are controlled include Cotton bollworm (Helicoverpa armigera), Codling moth (Cydia pomonella), Lightbrown apple moth (Epiphyas postvittana), and European corn borer (Ostrinia nubilalis).

Trichogramma species vary in their host specificity. This can lead to non-target hosts being parasitized. This can cause problems by reducing the amount of parasitism of the target host, and depending on the rate of parasitism, non-target effects could be significant on non-target host populations.

Species used

The most commonly used species for biological control are T. atopovirilia, T. brevicapillum, T. deion, T. exiguum, T. fuentesi, T. minutum, T. nubilale, T. platneri, T. pretiosum and T. thalense.[3]

Trichogramma pretiosum

Trichogramma pretiosum is the most widely distributed Trichogramma species in North America.[3] It has been the focus of many research studies and has been able to be reared on 18 genera of Lepidoptera. It is a more generalized parasitoid, able to parasitise a range of different species.

T. pretiosum was introduced into Australia in the 1970s as part of the Ord River Irrigation Area (ORIA) IPM scheme.[21][22]

Trichogramma carverae

Trichogramma carverae are mainly used for light brown apple moth and codling moth control and is predominately used in orchards.[23] In Australia T. carverae is used for biological control of light brown apple moth in vineyards. Though Australia has its own native Trichogramma species there has not been much work undertaken to commercially use them for biological control within Australia.[24]

Light brown apple moth is common throughout Australia and is polyphagous on more than 80 native and introduced species. The larvae are the stage that causes the most damage, especially to grape berries which provides sites for bunch rot to occur.[25] Losses in the crops can amount up to $2000/ha in one season. It is very predominant in areas like the Yarra Valley. Insecticide use is not a preferred method by most growers who prefer a more natural mean of controlling pests. As a result, Trichogramma were considered a good candidate for biological control as the larvae are difficult to control with insecticide and light brown apple moths are relatively vulnerable to egg parasitism with their eggs being laid in masses of 20-50 on the upper surfaces of basal leaves in grapevines.

Selected Species

References

  1. ↑ Flanders, S; Quednau, W (1960). "Taxonomy of the genus Trichogramma (Hymenoptera, Chalcidoidea, Trichogrammatidae)". BioControl. 5: 285–294. doi:10.1007/bf02372951.
  2. ↑ Consoli FL, Parra JRP, Zucchi RA (2010) 'Egg Parasitoids in Agroecosystems with Emphasis on Trichogramma.' (Springer).
  3. 1 2 3 4 5 Knutson A (2005) 'The Trichogramma Manual: A guide to the use of Trichogramma for Biological Control with Special Reference to Augmentative Releases for Control of Bollworm and Budworm in Cotton.' (Texas Agricultural Extension Service).
  4. ↑ Sumer, F; Tuncbilek, AS; Oztemiz, S; Pintureau, B; Rugman-Jones, P; Stouthamer, R (2009). "A molecular key to the common species of Trichogramma of the Mediterranean region". BioControl. 54: 617–624. doi:10.1007/s10526-009-9219-8.
  5. ↑ Upadhyay RK, Mukerji KG, Chamola BP (2001) 'Biocontrol potential and its Exploitation in Sustainable Agriculture: Insect Pests.' (Kluwer Academic/ Plenum Publishers).
  6. 1 2 Knutson A (2005) 'The Trichogramma Manual: A guide to the use of Trichogramma for Biological Control with Special Reference to Augmentative Releases for Control of bollworm and Budworm in Cotton.' (Texas Agricultural Extension Service).
  7. ↑ Macademia Nut Plantation, Trichogramma
  8. ↑ Klomp, H; Teerink, B.J.; Wei, Chun Ma (1979). "Discrimination Between Parasitized and Unparasitized Hosts in the Egg Parasite Trichogramma embryophagum (Hym.=Trichogrammatidae)=a Matter of Learning and Forgetting". Netherlands Journal of Zoology. Koninklijke Brill NV. 30 (2): 254–27. doi:10.1163/002829679X00412. ISSN 0028-2960. Retrieved May 22, 2014.
  9. ↑ Nagarkatti, S; Nagaraja, H (1977). "Biosystematics of Trichogramma and Trichogrammatoidea species". Annual Review of Entomology. 22: 157–176. doi:10.1146/annurev.en.22.010177.001105.
  10. ↑ Thomson, LJ; Rundle, BJ; Carew, ME; Hoffmann, AA (2003). "Identification and characterization of Trichogramma species from south-eastern Australia using the internal transcribed spacer 2 (ITS-2) region of the ribosomal gene complex". Entomologia Experimentalis et Applicata. 106: 235–240. doi:10.1046/j.1570-7458.2003.00029.x.
  11. ↑ Nagarkatti, S; Nagaraja, H (1971). "Redescriptions of some known species of Trichogramma (Hym., Trichogrammatidae), showing the importance of the male genitalia as a diagnostic character". Bulletin of Entomological Research. 61: 13–31. doi:10.1017/s0007485300057412.
  12. ↑ Polaszek, A; Rugman-Jones, P; Stouthamer, R; Hernandez-Suarez, E; Cabello, T; Pino Pérez, M (2012). "Molecular and morphological diagnoses of five species of Trichogramma: biological control agents of Chrysodeixis chalcites (Lepidoptera: Noctuidae) and Tuta absoluta (Lepidoptera: Gelechiidae) in the Canary Islands". BioControl. 57: 21–35. doi:10.1007/s10526-011-9361-y.
  13. ↑
  14. 1 2 3 Grenier, Simon; et al. (1998). "Successful horizontal transfer of Wolbachia symbionts between Trichogramma wasps". Proceedings of the Royal Society of London B: Biological Sciences. 265: 1441–1445. doi:10.1098/rspb.1998.0455.
  15. 1 2 3 4 5 6 Huigens, M. E.; et al. (2004). "Natural interspecific and intraspecific horizontal transfer of parthenogenesis–inducing wolbachia in trichogramma wasps". Proceedings of the Royal Society of London B: Biological Sciences. 271: 509–515. doi:10.1098/rspb.2003.2640.
  16. 1 2 Bourtzis, Kostas; O'Neill, Scott (1998). "Wolbachia Infections and Arthropod Reproduction". BioScience. 48: 287–293. doi:10.2307/1313355.
  17. 1 2 3 4 Charlat, Sylvain; Hurst, Gregory D. D.; Merçot, Hervé (2003-04-01). "Evolutionary consequences of Wolbachia infections". Trends in genetics: TIG. 19 (4): 217–223. doi:10.1016/S0168-9525(03)00024-6. ISSN 0168-9525. PMID 12683975.
  18. ↑ Smith SM (1996) Biological control with Trichogramma: advances, successes, and potential of their use. In 'Annual Review of Entomology' pp. 375-406.
  19. ↑ Flanders, SE (1930). "Mass Production of Egg Parasites of the Genus Trichogramma". Hilgardia. 4: 465–501. doi:10.3733/hilg.v04n16p465.
  20. ↑ Hassan, SA (1993). "The mass rearing and utilization of Trichogramma to control lepidopterous pests: Achievements and outlook". Pesticide Science. 37: 387–391. doi:10.1002/ps.2780370412.
  21. ↑ Davies, AP; Zalucki, MP (2008). "Collection of Trichogramma Westwood (Hymenoptera: Trichogrammatidae) from tropical northern Australia: a survey of egg parasitoids for potential pest insect biological control in regions of proposed agricultural expansion". Australian Journal of Entomology. 47: 160–167. doi:10.1111/j.1440-6055.2008.00644.x.
  22. ↑ Davies, AP; Pufke, US; Zalucki, MP (2011). "Spatio-temporal variation in Helicoverpa egg parasitism by Trichogramma in a tropical Bt-transgenic cotton landscape". Agricultural and Forest Entomology. 13: 247–258. doi:10.1111/j.1461-9563.2010.00512.x.
  23. ↑ Llewellyn R (2002) The good bug book: beneficial organisms commercially available in Australia and New Zealand for biological pest control.' (Integrated Pest Management Pty Ltd).
  24. ↑ Glenn, DC; Hercus, MJ; Hoffmann, AA (1997). "Characterizing Trichogramma (Hymenoptera: Trichogrammatidae) species for biocontrol of light brown apple moth (Lepidoptera: Tortricidae) in grapevines in Australia". Annals of the Entomological Society of America. 90: 128–137.
  25. ↑ Glenn, DC; Hoffmann, AA (1997). "Developing a commercially viable system for biological control of light brown apple moth (Lepidoptera: Tortricidae) in grapes using endemic Trichogramma (Hymenoptera: Trichogrammatidae)". Journal of Economic Entomology. 90: 370–382.
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