Inversion genotype and biological characteristics of
In India malaria endemicity is characterized by diverse ecology and multiple disease vector species . In the Southeast Asian region, India alone contributes to nearly 80% of malaria cases with the largest population of the world living at risk of malaria. In 2011, India reported 1.3 million confirmed malaria cases and 753 attributable deaths, but estimated cases and deaths are 10 to 20 times more [2,3]. Of the two
Mosquito fauna is rich in the tropical climate with numerous and diverse breeding resources . Of 58 anophelines in India, only six taxa are major malaria vectors with regional distribution (Figure 1).
India is experiencing rapid ecological changes owing to population explosion, urbanization, development projects, deforestation and human migration affecting mosquito ecology and disease transmission. In the recent past, significant progress has been made in understanding the genetics and bionomics of the disease vectors, and in the development of newer control tools to strengthen primary healthcare services specific to India [9-14]. In this chapter we shall restrict systematic review on dominant
Anopheles (Cellia) culicifacies Giles species complex
Sibling species were initially characterized by species specific diagnostic fixed paracentric inversions readable in polytene chromosomes suggestive of pre-mating barriers in field populations [18-24], and further substantiated by number of techniques including post-zygotic isolation mechanisms in laboratory conditions , mitotic karyotype Y- chromosome polymorphism [26-28], gene enzyme variation , cuticular hydrocarbon profiles , and species specific DNA probes . Recently, PCR-based diagnostic assays were developed for sequencing 28S-D3 domain , ITS2-PCR-RFLP , rDNA ITS2 region , which grouped
|Inversion genotype||X+a+b; 2+g1+h1; +i1/i1||Xab; 2g1+h1||Xab; 2+g1h1||X+a+b; 2i1+h1||Xab; 2g1+h1;|
|Anthropophilic Index (%)||0-4||0-1||0-3||0-1||80|
(Peak biting activity)
|Sporozoite infection rate (%)||0.51||0.04||0.3||0.4||20|
|Breeding preferences||Rainwater, clean irrigation water||Riverine ecology||Rainwater, clean irrigation water||Rainwater, clean irrigation water||Riverine ecology|
|Rate of development of resistance|
|DDT||Slow (9-10 yr)||Fast
|No data||No data|
|Malathion||Slow (9-10 yr)||Medium
|No data||No data|
|No data||No data|
The distribution, relative abundance and predominance of sibling species (but not exclusive) is given in Figure 2. Among its sibling species, species B is the most predominant throughout the country and occurs sympatrically in most areas with predominance of species A in the north and species B in the south . In eastern Uttar Pradesh, north Bihar and northeastern states, species B is either predominant or the only prevalent species. Species B and C are sympatric in western and eastern India. Species D is sympatric with species A and B in northwestern region, and with species A, B and C in central southern India. Species E is sympatric with species B in southern Tamilnadu including Rameshwaram islands. The proportions of sibling species, however, varied in different geographical zones and seasons, e.g., in Alwar (state of Rajasthan), species B proportions increased in post-monsoon months; whereas proportions of species D remained the same throughout the year and density of species C remained very low .
All member sibling species of the
However, more information on distribution and bionomics of species E is deemed necessary to substantiate its distribution range and role in malaria transmission in India. In addition, understanding population structure of
Anopheles (Cellia) fluviatilis James species complex
|S||+q’+r’||Low to Moderate (1-40)||Anthropophilic||Human dwellings||Hilly forests & foothills||Hyperendemic|
|T||q’+r’||High (up to 200)||Almost totally zoophilic||Cattle sheds||Foothills & plains||Hypo - mesoendemic|
Sibling species S is highly anthropophilic and responsible for maintaining hyperendemic malaria predominantly in state of Odisha (formerly Orissa), eastern India . It prefers to rest indoor human dwellings and have been incriminated and proven to be an efficient vector in areas of its distribution [61,62]. Sibling species T is widely distributed but is largely zoophilic and rests in cattle sheds . Sibling U holds similar characteristics but has limited distribution range presently restricted to northern India. Chen et al  documented three haplotypes in species T (designated T1, T2, Y) with its distribution in India, Nepal, Pakistan and Iran implicating the existence of additional taxa within the An
Preferred breeding habitats are seepage water streams with perceptible flow of water, river margins, irrigation channels, shallow wells, terraced rice fields along foothills etc [7,11,65]. Peak biting activity occurs between 20:00 to 24:00 hours but it may vary in different seasons and locations. Both
For control of
Anopheles (Cellia) minimus Theobald species complex
Consequently, systematic studies by independent investigators revealed the reappearance of
Ever since initial recognition of
Based on DNA sequences of internal transcribed spacer 2 (ITS2) and D3 domain of 28S rDNA (28S-D3) of morphologically identified
It is suggested that in areas with
Anopheles (Cellia) dirus Peyton & Harrison species complex
Among these member species, only
Even though populations of
Anopheles (Cellia) sundaicus (Rodenwaldt) species complex
In Andaman and Nicobar islands,
Anopheles (Cellia) stephensi Liston – A complex of variants
The species is resistant to multiple insecticides but indoor residual spraying is not used for control. Instead recommended control measures are (i) source reduction, (ii) minor engineering interventions (iii) anti-larval methods including chemical and biological larvicides, (iv) application of larvivorous fish, i.e., guppy and gambusia, (v) aerosol space spraying for control of adult vector populations, (vi) legislative bylaws for preventing mosquito breeding . In the face of rapid urbanization, unplanned growth and mushrooming of urban slums, rationed water supply and unsafe water storage practices; urban malaria is a growing problem presently accounting for >10% reported malaria cases in the country . Overall, malaria cases in the rural and urban areas are grossly underestimated due to scanty surveillance and unreliable laboratory services.
8. Prospects of vector control and research priorities
India has about a billion population at risk of malaria and accounts for the highest disease burden in Southeast Asia for estimated loss of disability adjusted life years [3,6]. Malaria transmission is complex due to multi-species co-existence and variable species dominance and bionomical characteristics [13,14]. Although, transmission trends seem to be declining (Figure 9), National Vector Borne Disease Vector Control Programme (NVBDCP) is faced with new emerging challenges. Some of these are (i) multiple insecticide resistance against target disease vector mosquito species, (ii) emerging multi-drug resistance and steadily rising proportions of
Indoor residual spraying (IRS) for vector control has become less effective and operationally difficult proposition [9,94]. In addition, ecological driven changes, population migration across borders, deforestation, developmental projects, and poor infrastructure have led to the opportunities for vector proliferation and increased malaria receptivity. Due to poor community acceptance for IRS and spray coverage of target population groups , India has embarked upon large scale implementation of Insecticide-treated netting materials / long-lasting insecticidal nets (LLINs) prioritizing high-risk population in malaria endemic states/districts. Disease transmission trends are declining in beneficiary population groups (formerly intractable high-risk areas); hence it is the right time to siege the opportunity for up scaling LLIN based intervention coupled with appropriate drug policy in place to combat the malaria illness and preventing spread of drug-resistant malaria [112,113,168-170]. It is worrisome, however, that the LLINs presently in use employ only pyrethroids, and
Besides dominant proven vector species, sporadic gut/ gland infections have also been recorded in
In moving forward for achieving ambitious goal of malaria elimination in feasible districts/states, lot more needs to be accomplished in understanding vector bionomics in the altered ecology. The future priority area should include developing malaria-risk maps for focused interventions, ecological succession of disease vector species, monitoring insecticide resistance, cross-border initiative with neighboring countries for data sharing and coordinated control efforts, development of evidence-based newer tools for vector control, strengthening health systems for improved surveillance and monitoring, and universal access to malaria treatment and prevention which would help meeting the Millennium Development Goal in reducing malaria morbidity and mortality by 2015 [191-193].
During the past decade, there has been significant progress in development of molecular techniques in identification of sibling species of the dominant mosquito vector taxa, understanding their bionomical characteristics and role in malaria transmission in India. Among these, for
We are thankful to Drs. T. Adak, K. Raghavendra, O.P. Singh, N. Nanda, A. Das, A. Kumar, S.K. Ghosh for access to the valued literature and consultations. We are also indebted to Dr. S. Manguin for encouragement and advice for development of the manuscript. This submission has been approved by the Institute Publication Screening Committee and bears the approval No. 022/2012.
DDT: Dichloro-diphenyl-trichloroethane; rDNA: Ribosomal deoxyribonucleic acid; ITS2: Internal Transcribed Spacer 2; PCR: Polymerase Chain Reaction; RFLP: Restricted Fragment Length Polymorphism; CO II: Cytochrome Oxidase II; IRS: Indoor Residual Spray; LLIN: Long-lasting Insecticidal Net; MPO: Modified Plan of Operation; NVBDCP: National Vector Borne Disease Control Programme.
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