Anopheles gambiae

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Anopheles gambiae
AnophelesGambiaemosquito.jpg
Scientific classification
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Species complex:
A. gambiae sensu lato
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Binomial name
Anopheles gambiae
The tube-like heart (green) extends along the body, interlinked with the diamond-shaped alary muscles (also green) and surrounded by pericardial cells (red). Blue depicts cell nuclei. Structural organization of the heart of the mosquito Anopheles gambiae - image.ppat.v08.i11.g001.png
The tube-like heart (green) extends along the body, interlinked with the diamond-shaped alary muscles (also green) and surrounded by pericardial cells (red). Blue depicts cell nuclei.

The Anopheles gambiae complex consists of at least seven morphologically indistinguishable species of mosquitoes in the genus Anopheles . The complex was recognised in the 1960s and includes the most important vectors of malaria in sub-Saharan Africa, particularly of the most dangerous malaria parasite, Plasmodium falciparum . [2] It is one of the most efficient malaria vectors known. The An. gambiae mosquito additionally transmits Wuchereria bancrofti which causes lymphatic filariasis, a symptom of which is elephantiasis. [3]

Contents

Discovery and elements

The Anopheles gambiae complex or Anopheles gambiae sensu lato was recognized as a species complex only in the 1960s. The A. gambiae complex consists of:

The individual species of the complex are morphologically difficult to distinguish from each other, although it is possible for larvae and adult females. The species exhibit different behavioural traits. For example, Anopheles quadriannulatus is both a saltwater and mineralwater species. A. melas and A. merus are saltwater species, while the remainder are freshwater species. [7] Anopheles quadriannulatus generally takes its blood meal from animals (zoophilic), whereas Anopheles gambiae sensu stricto generally feeds on humans, i.e. is considered anthropophilic.[ citation needed ] Identification to the individual species level using the molecular methods of Scott et al. (1993) [8] can have important implications in subsequent control measures.

Anopheles gambiae in the strict sense

An. gambiae sensu stricto (s.s.) has been discovered to be currently in a state of diverging into two different species—the Mopti (M) and Savannah (S) strains—though as of 2007, the two strains are still considered to be a single species. [9] [10]

A mechanism of species recognition using the sound emitted by the wings and identified by Johnston's organ was proposed in 2010, [11] however this mechanism has never been confirmed since, and the overall mechanism theory through "harmonic convergence" has been challenged. [12] [13]

Genome

An. gambiae s.s. genomes have been sequenced three times, once for the M strain, once for the S strain, and once for a hybrid strain. [9] [10] Currently, ~90 miRNA have been predicted in the literature (38 miRNA officially listed in miRBase) for An. gambiae s.s. based upon conserved sequences to miRNA found in Drosophila .[ citation needed ] Holt et al., 2002 and Neafsey et al., 2016 find transposable elements to be ~13% of the genome, similar to Drosophila melanogaster (also in Diptera). [14] However they find the proportion of TE types to be very different from D. melanogaster with approximately the same composition of long terminal repeat retrotransposons, non-long terminal repeat retrotransposons and DNA transposons. [14] These proportions are believed to be representative of the genus. [14]

The genetics and genomics of sex chromosomes have been discovered and studied by Windbichler et al., 2007 and Galizi et al., 2014 (a Physarum polycephalum homing endonuclease which destroys X chromosomes), Windbichler et al., 2008 and Hammond et al., 2016 (methods to reduce the female population), Windbichler et al., 2011 (trans from yeast), Bernardini et al., 2014 (a method to increase the male population), Kyrou et al., 2018 (a female necessary exon and a homing endonuclease to drive it), Taxiarchi et al., 2019 (sex chromosome dynamics in general) and Simoni et al., 2020 (an X chromosome destroying site specific nuclease). [15] See § Gene drive below for their applications.

An. gambiae has a high degree of polymorphism. This is especially true in the cytochrome P450s, Wilding et al., 2009 finding 1 single nucleotide polymorphism (SNP)/26 base pairs. This species has the highest amount of polymorphism in the CYPs of any insect known, much tending to be found in "scaffolds" that are found only in particular subpopulations. These are termed "dual haplotype regions" by Holt et al., 2002 who sequenced the PEST strain. [16] :241

In common with many chromosomes, An. gambiae codes for spindle and kinetochore-associated proteins. Hanisch et al., 2006 locate AgSka1 , the spindle and kinetochore-associated protein 1 gene, at EAL39257. [17]

The entire Culicidae family may or may not conserve epigenetic mechanisms as of 2012 this remains unresolved. [18] Toward answering this question, Marhold et al., 2004 compare their own previous work in Drosophila melanogaster against new sequences of D. pseudoobscura and An. gambiae. [18] They find all three do share the DNA methylation enzyme DNMT2 ( DmDNMT2 , DpDNMT2 , and AgDNMT2). [18] This suggests all Diptera may conserve an epigenetic system employing Dnmt2. [18]

Hosts

Hosts include Bos taurus , Capra hircus , Ovis aries and Sus scrofa . [19]

Parasites

Parasites include Plasmodium berghei (for which it also serves as a vector), [20] [21] [22] and the bioinsecticides/entomopathogenic fungi Metarhizium robertsii [20] and Beauveria bassiana . [20] All three of these parasites combine with insecticides to reduce fitness see § Insecticides below. [20] CRISPR/Cas9 and U6-gRNA are increasingly (as of 2020) being used together for knockout experiments in mosquitoes. [22] Dong et al., 2018 develops and presents a new U6-gRNA+Cas9 technique in An. gambiae, and utilizes it to knock out fibrinogen related protein 1 (FREP1), thereby severely reducing infection of the mosquito by P. berghei and P. falciparum . [22] However this also demonstrates the centrality of FREP1 to the insect's success, impairing all measured activities across all life stages. [22] Yang et al., 2020 uses the Dong method to do the same with mosGILT , also severely reducing Plasmodium infection of the mosquito but also finding a vital life process is impaired, in mosGILT's case ovary development. [22]

Control

Insecticides

Parasites/bioinsecticides and chemical insecticides synergistically reduce fitness. Saddler et al., 2015 finds even An. gambiae with knockdown resistance (kdr) are more susceptible to DDT if they are first infected with Plasmodium berghei [20] [21] and Farenhorst et al., 2009 the same for Metarhizium robertsii or Beauveria bassiana . [20] This is probably due to an effect found by Félix et al., 2010 and Stevenson et al., 2011: An. gambiae alters various activities especially CYP6M2 in response to P. berghei invasion. CYP6M2 is known to somehow produce pyrethroid resistance, and pyrethroids and DDT share a mechanism of action. [21]

Gene drive

Research relevant to the development of gene drive controls of An. gambiae have been performed by Windbichler et al., 2007, Windbichler et al., 2008, Windbichler et al., 2011, Bernardini et al., 2014, Galizi et al., 2014, Hammond et al., 2016, Kyrou et al., 2018, Taxiarchi et al., 2019 and Simoni et al., 2020. [15] For specific genes involved see § Genome above. These can all be used in pest control because they induce infertility. [15]

Fecundity

Fecundity of An. gambiae depends on the detoxification of reactive oxygen species (ROS) by catalase. [23] Reduction in catalase activity significantly reduces reproductive output of female mosquitoes, indicating that catalase plays a central role in protecting oocytes and early embryos from ROS damage. [23]

Historical note

An. gambiae invaded northeastern Brazil in 1930, which led to a malaria epidemic in 1938/1939. [24] The Brazilian government assisted by the Rockefeller Foundation in a programme spearheaded by Fred Soper eradicated these mosquitoes from this area. This effort was modeled on the earlier success in eradication of Aedes aegypti as part of the yellow fever control program. The exact species involved in this epidemic has been identified as An. arabiensis . [25]

Peptide hormones

Kaufmann and Brown 2008 find the An. gambiae adipokinetic hormone (AKH) mobilizes carbohydrates but not lipids. Meanwhile AKH/Corazonin Peptide (ACP) does not mobilize (or inhibit mobilization) of either. Mugumbate et al., 2013 provides in solution and membrane bound structures from a nuclear magnetic resonance investigation. [26]

Related Research Articles

<span class="mw-page-title-main">Malaria</span> Mosquito-borne infectious disease

Malaria is a mosquito-borne infectious disease that affects vertebrates. Human malaria causes symptoms that typically include fever, fatigue, vomiting, and headaches. In severe cases, it can cause jaundice, seizures, coma, or death. Symptoms usually begin 10 to 15 days after being bitten by an infected Anopheles mosquito. If not properly treated, people may have recurrences of the disease months later. In those who have recently survived an infection, reinfection usually causes milder symptoms. This partial resistance disappears over months to years if the person has no continuing exposure to malaria.

<i>Anopheles</i> Genus of mosquito

Anopheles is a genus of mosquito first described by the German entomologist J. W. Meigen in 1818, and are known as nail mosquitoes and marsh mosquitoes. Many such mosquitoes are vectors of the parasite Plasmodium, a genus of protozoans that cause malaria in birds, reptiles, and mammals, including humans. The Anopheles gambiae mosquito is the best-known species of marsh mosquito that transmits the Plasmodium falciparum, which is a malarial parasite deadly to human beings; no other mosquito genus is a vector of human malaria.

<i>Wolbachia</i> Genus of bacteria in the Alphaproteobacteria class

Wolbachia is a genus of gram-negative bacteria that can either infect many species of arthropod as an intracellular parasite, or act as a mutualistic microbe in filarial nematodes. It is one of the most common parasitic microbes of arthropods, and is possibly the most common reproductive parasite in the biosphere. Its interactions with its hosts are often complex. Some host species cannot reproduce, or even survive, without Wolbachia colonisation. One study concluded that more than 16% of neotropical insect species carry bacteria of this genus, and as many as 25 to 70% of all insect species are estimated to be potential hosts.

<span class="mw-page-title-main">Microsporidia</span> Phylum of fungi

Microsporidia are a group of spore-forming unicellular parasites. These spores contain an extrusion apparatus that has a coiled polar tube ending in an anchoring disc at the apical part of the spore. They were once considered protozoans or protists, but are now known to be fungi, or a sister group to true fungi. These fungal microbes are obligate eukaryotic parasites that use a unique mechanism to infect host cells. They have recently been discovered in a 2017 Cornell study to infect Coleoptera on a large scale. So far, about 1500 of the probably more than one million species are named. Microsporidia are restricted to animal hosts, and all major groups of animals host microsporidia. Most infect insects, but they are also responsible for common diseases of crustaceans and fish. The named species of microsporidia usually infect one host species or a group of closely related taxa. Approximately 10 percent of the known species are parasites of vertebrates — several species, most of which are opportunistic, can infect humans, in whom they can cause microsporidiosis.

<span class="mw-page-title-main">Mosquito net</span> Fine net used to exclude mosquitos and other biting insects

A mosquito net is a type of meshed curtain that is circumferentially draped over a bed or a sleeping area, to offer the sleeper barrier protection against bites and stings from mosquitos, flies, and other pest insects, and thus against the diseases they may carry. Examples of such preventable insect-borne diseases include malaria, dengue fever, yellow fever, zika virus, Chagas disease and various forms of encephalitis, including the West Nile virus.

<i>Plasmodium berghei</i> Single celled parasite, rodent malaria

Plasmodium berghei is a single-celled parasite causing rodent malaria. It is in the Plasmodium subgenus Vinckeia.

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Plasmodium yoelii is a parasite of the genus Plasmodium subgenus Vinckeia. As in all Plasmodium species, P. yoelii has both vertebrate and insect hosts. The vertebrate hosts for this parasite are mammals.

<i>Anopheles culicifacies</i> Species of mosquito

Anopheles culicifacies is a mosquito species complex and one of the primary vectors of malaria on the Indian subcontinent. It consists of five sibling species, provisionally designated as species A, B, C, D, and E. It prefers to rest indoors in cattle sheds, where it feeds on cattle. The control of A. culicifacies has become difficult due to the development of insecticide resistance against all commonly used insecticides, including new-generation insecticides such as synthetic pyrethroids.

VectorBase is one of the five Bioinformatics Resource Centers (BRC) funded by the National Institute of Allergy and Infectious Diseases (NIAID), a component of the National Institutes of Health (NIH), which is an agency of the United States Department of Health and Human Services. VectorBase is focused on invertebrate vectors of human pathogens working with the sequencing centers and the research community to curate vector genomes.

<i>Anopheles stephensi</i> Species of fly

Anopheles stephensi is a primary mosquito vector of malaria in urban India and is included in the same subgenus as Anopheles gambiae, the primary malaria vector in Africa. A. gambiae consists of a complex of morphologically identical species of mosquitoes, along with all other major malaria vectors; however, A. stephensi has not yet been included in any of these complexes. Nevertheless, two races of A. stephensi exist based on differences in egg dimensions and the number of ridges on the eggs; A. s. stephensisensu stricto, the type form, is a competent malaria vector that is found in urban areas, and A. s. mysorensis, the variety form, exists in rural areas and exhibits considerable zoophilic behaviour, making it a poor malaria vector. However, A. s. mysorensis is a detrimental vector in Iran. An intermediate form also exists in rural communities and peri-urban areas, though its vector status is unknown. About 12% of malaria cases in India are due to A. stephensi.

Thioester containing protein 1, often called TEP1 is a key component of the arthropod innate immune system. TEP1 was first identified as a key immunity gene in 2001 through functional studies on Anopheles gambiae mosquitoes.

<i>Anopheles sinensis</i> Species of mosquito

Anopheles sinensis is a species of mosquito that transmits malaria as well as lymphatic filariasis. It is regarded as the most important vector of these human parasitic diseases in Southeast Asia. It is the primary vector of vivax malaria in many regions. In China it also transmits the filalarial parasite, and arthropod roundworm. In Japan it is also a vector of a roundworm Setaria digitata in sheep and goats.

<span class="mw-page-title-main">Thorselliaceae</span> Genus of bacteria

Thorselliaceae is a family of bacteria belonging to the class Gammaproteobacteria and it was first described in February 2015. It is not assigned to an order. The family consists of four species in two genera. The bacteria are Gram-negative and rod shaped, approximately 1 μm wide and 2 μm long. They are facultative anaerobes and motile. Thorselliaceae bacteria have been found around the world associated with vector mosquitoes, mainly with vectors of malaria.

Anopheles nili is a species of mosquito in the Culicidae family. It comprises the following elements: An. carnevalei, An. nili, An. ovengensis and An. somalicus. The scientific name of this species was first published in 1904 by Theobald. It is the main mosquito species found in the south Cameroon forest zone which bites humans. It is known as a problematic carrier of malaria, although newly discovered, closely related species in the same genus have also been found to interact with A. nili as a disease vector. In that, they both have similar feeding habits on local targets in the Cameroon region.

<span class="mw-page-title-main">Flaminia Catteruccia</span> Italian professor of immunology and infectious disease

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<i>Anopheles arabiensis</i> African mosquito and disease vector

Anopheles arabiensis is a zoophilic species of mosquito and a vector of disease endemic to Africa.

Nora J. Besansky is an American molecular biologist. She is the Martin J. Gillen Professor of Biological Sciences at the University of Notre Dame. In 2020, Besansky was elected a Member of the National Academy of Sciences for being an expert in the genomics of malaria vectors.

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