Chironomids (Diptera) as Model Organisms An Appraisal

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CHIRONOMIDS (DIPTERA) AS MODEL ORGANISMS: AN APPRAISAL
Kaushik Sanyal, Atrayee Deya, Niladri Hazra
Entomology Research Unit, Department of Zoology, The University of Burdwan,
Burdwan 713104, West Bengal India
Corresponding author : [email protected]
ABSTRACT
The Chironomids (Diptera: Chironomidae), commonly known as “non-biting midges” are abundant, diverse
and predominantly widespread aquatic insects. They have high species richness due to their adaptive nature
in diverse habitats, ranging from fresh water to brackish or marine water. In recent years, interest in the food
and feeding behaviour of larval chironomids has expanded due to energetic importance in freshwater ecosystems,
improvement of taxonomic keys and controlling pestiferous emergence. A variety of factors including larval size,
food quality and sediment composition can influence the feeding behaviour of larval midges. They form an
integral link in aquatic food chains, constituting a biological connection between aquatic and terrestrial life.
Increasing eutrophication of aquatic ecosystem due to natural and anthropogenic factors resulting in rapid
colonization of these midges has enabled them to introduce as good bioindicator of water quality. Both larvae
and adult midges have been reported to cause human allergy and rice and glasshouse pests for its damaging
nature of field crops. The insects have been used as model organism in various aspects of cytogenetics,
karyosystematics, cytochemistry and gene amplification and in the study of phylogeny and biogeography. They
are essential components of multidisciplinary, palaeolimnological reconstruction of past, climate change, and
have been used to trace palaeoproductivity of lake systems, to reconstruct the history of acidification in lakes
to assess anthropogenic eutrophication and to monitor the impact of salinity fluctuations.
Key Words. Chironomidae; model organisms; bio-indicator; biogeography, vicariance, dispersal, palaeolimnology,
Introduction
Chironomids are widely distributed and constitute highly diversified group of aquatic holometab-
olous insects frequently occurring in high density in all kinds of limnic ecosystem (Coffman and
Ferrington 1996). Although the impact of these midges excludes human-feeding and disease transmis-
sion, they are still the subject of much importance. Interest in the food and feeding behaviour of larval
chironomids has been expanded in recent years due to energetic importance in freshwater ecosystems,
improvement of taxonomic keys and controlling pestiferous emergence. The larvae form a biological
connection between aquatic and terrestrial habitat (Mason 1973) and humans are ultimately linked to
these systems (Warwick 1990). They are known to play prominent role in biological systems of
classifying lakes (Brundin 1949) and are proving usefulness in palaeoecological studies of lake
sediments (Brodin and Gransborg 1993; Warwick 1980; Walker et al. 1991). At the suborganismal
level, the presence of giant polytene chromosomes in the larval salivary glands has encountered much
molecular and cytological research. Increasing demand for chironomid larvae in the aquaculture sector
forces the researchers to study its production, behaviour, development, growth, feeding, and nutritive
value. The dominance of the larvae in aquatic ecosystems means that they are unavoidably
encountered by every freshwater ecologist. Adult flies can impinge on our activities by their nuisance
numbers outdoors or through their ability to thrive in domestic water distribution systems. Increasing
aquatic eutrophication due to natural and anthropogenic factors seems conducive for the rapid
colonization of the midges throughout the globe. Moreover, the chironomids integrate various biological
ISBN 978-81-927762-0-0
Proc. Nat. Con. Cha. Bio. Res. Man.

189
processes and may be used as indicators of environmental conditions (Kuhlmann et al. 2001) for its
relatively long life cycle and low mobility.
Habit and Habitat
The midges are adapted to varied habitats, the range of which is unparalleled among other insect
groups. Majority of them occur in freshwater (Fig. 1) but species are also found marine, brackish
water and terrestrial habitats. Their abode include the littoral and benthic regions of lakes, ponds,
temporary pools, temporary to permanent ditches, shallow stagnant waters, rice fields, puddles, seeps,
seasonally flooded tree–holes, bromeliads, leaf moulds, hollow stumps of bamboo, water filled axils of
plants, tree holes, waterfalls, glacial melt-water, hot and cold springs, mountain streams, fast and slow–
flowing rivers and marine littorals (Ashe 1983).
Feeding Behavior
It is known that the feeding habits of midge larvae are rather varied. Generally, these larvae ingest
five kinds of food such as algae, detritus and associated microorganisms, macrophytes, wood debris,
and invertebrates (Berg 1995). Based upon the feeding mode, larvae can be grouped in six categories:
collectors (gatherers and filterers), shredders, scrapers, and predators (engulfers and piercers)
(Coffman and Ferrington 1996). According to Pinder (1992), the larvae fed with food rich in diatoms
were found to survive better with quick growth and development than those provided with food of
predominant fine organic detritus.
Chironomid as Food
The worms are considered as a natural diet for fish, dabbling or diving ducks, and water fowl and
other invertebrates (Willassen and Bjørklund 1989; Wrubleski and Rosenberg 1990). Armitage (1995)
has suggested that the preference of bottom feeder organisms for chironomid larvae and pupae as food
source is related to their high energy content (4.1–6.1 KCal g-1). They can also be used as frozen
food when added to shrimp meal for the freshwater prawn, Macrobrachium rosenbergii, resulting
improvement in growth rate and survival under laboratory conditions (Abdel- Razek et al. 1998).
Growing demand for chironomid larvae in the aquaculture sector compels the researchers to study and
enhance its production, behaviour, development, growth, feeding and nutritive value. Midge larvae are
excellent source of protein (Noue and Choubert 1985) and other nutrients, such as, lipid, vitamins and
minerals (Mclarney et al. 1974). In aquaculture practices, minerals might be supplemented to fish and
shrimps by feeding them with chironomid larvae (Habib et al. 1997). The protein content of chironomid
larvae has been estimated as 56% (Sugden 1973). Chironomid larvae possess high digestibility (73.6%)
value and the apparent function in small quantities as a growth promoter in fish, crustaceans, larvae
etc. It has therefore, become imperative to make a suitable culture system for the chironomid larvae
(Noue and Choubert 1985).
Biogeographical Analyses Aspect
The Chironomid midges are ubiquitous, and represent the most abundant macroinvertebrate group
in numbers and diversity of species prevailing in all zoo–geographical regions (Ashe 1983). Globally,
there are estimated, 20,000 species of which 4,000 species presently known (Fig. 2) (Ferrington Jr.
2008). Different species thrive in different environments; some have widespread ecological tolerances
whereas some are more restricted. The diversity and distribution of Chironomidae makes it well suited
for geographic co-evolutionary analyses. They are used for study of distributional patterns, explanation
of such patterns like dispersal, vicariance etc. (Fig. 3) (Nelson and Platnick 1984), and centre of origin,
range extension and area of endemism. Dispersal and vicariance are often considered competing
hypotheses in historical biogeography. Disjunct distribution patterns can be explained either by

190
fragmentation of widespread ancestors by vicariant events or by dispersal across a preexisting barrier.
Current methods of biogeographic analyses are based on the vicariance model because nearly any
distribution pattern can be explained by dispersal, making dispersal hypotheses resilient to falsification
(Morrone and Crisci 1995). More stable phylogenies in the future will certainly add valuable
information to existing hypotheses on zoogeographical patterns (Sæther and Ekrem 2003).
Palaeolimnological Study Aspect
The midges are essential components of multidisciplinary, palaeolimnological reconstruction of past,
to reconstruct the history of acidification in lakes (Brodin and Gransborg 1993) and also in
palaeoclimatic study (Rosenberg 1993; Brooks 2006). Fossil chironomids from lake sediments are
frequently used as bioindicator in studies of paleoproductivity of lake system (Bryce 1962; Smol and
Douglas 2005). The direct or indirect responsiveness of chironomid fauna to the changes of water
quality and the long-term persistence of useful taxonomic characters associated with the head capsule
and mouthparts has led to the widespread use of their remains in palaeoecological studies of lakes.
Now-a-day, multivariate statistical techniques (DCA, CCA) have measured the correlation between
distribution of chironomids and surface water temperature (Walker et al. 1991). Because many of the
taxa have stenothermic distribution (Hofmann 1986), fossil chironomids appear to be good indicator of
water and air-temperature in many lake ecosystems. Studies of head capsules in late-glacial lake
sediments have shown that midges respond rapidly to climatic fluctuations (Lotter et al. 1998). Human
impacts, such as forestry, agriculture and atmospheric pollution, can impair our ability to quantify
chironomid-climate relationship, because community composition may be strongly affected by changes
in nutrients and food-supply, sediment composition and deepwater oxygen availability unrelated to
climatic conditions (Clerk et al. 2000).
Ecological Study Aspect
Chironomid larvae have been considered to be the most promising biological indicator of water
quality among fresh water macro-invertebrates because of their ubiquity and abundance in aquatic
ecosystems and also are essential components of quantitative and qualitative community approaches
to bio-monitoring. Chironomids species diversity and their sensitivity to eutrophic conditions have been
used to create trophic status classification of lakes (Oligotrophic, mesotrophic and eutrophic). The
larval stages of chironomid are shredders and thus contribute largely towards the recycling of the
detritus into available nutrients, besides serving as prey for different taxonomic group of predators.
Thus the chironomids play an important role in the aquatic food chain by forming aquatic food web
and make a major link between producers, such as phytoplankton, benthic algae and secondary
consumers (Tokeshi 1995). They are also broadly dispersed and often the most abundant insect species
in freshwater ecosystems (Sæther 1979) and their ecological diversity is proved by their physiological
tolerance to environmental stress. The midges in general are tolerant to heavy metal contamination
(Pinder 1986). Morphological deformities and levels of fluctuating asymmetry in organisms are caused
by several factors, such as developmental instabilities, environmental stress, and genetic factors
(Clarke 1995). The rate of mouthpart deformities varied considerably among populations and
generations during the multigenerational study (Vogt et al. 2013). Morphological features of the larvae
indicate toxic stress (Warwick 1990) (Figs. 4a-d). Studies of population dynamics show that
chironomids play an important role in preserving sediment quality. They are an essential element in the
organic matter circulation of the lake (Dévai, 1990).

191
Impacts of Chironomids
Both midge larvae and adults have positive and negative impacts as depicted in figure 5.
Discussion
Chironomid midges may be considered as good model organisms as they are not only used in the
study of systematics but also for its trends in ecology, palaeolimnology, biogeography, cytology and
ecotoxicological research. Chironomid midges are one of the good indicators of qualitative components
of aquatic ecosystems reflecting the nutrient status of lakes and streams (Rosenberg 1993). Different
species of the midges have been utilized in elucidation of hydro-geological features of earth
biomonitoring (Bonada et al. 2006) and in assessment of pollutant load of aquatic environment
(Bhattacharyay et al. 2005, 2006; Domingues et al. 2008). They are helpful in predicting the diversity
and strength of food web of aquatic communities (Bersier et al. 2002). Despite of its small size, they
have complex repertoire of behavioral responses including activities that relate readily to the
environmental condition. Apart from being a model organism, the midges are now-a-days utilized as
fish meal for commercial fish production.
Acknowledgement
We are grateful to Dr. P. K. Chaudhuri, Former Professor, Dept. of Zoology, The University of
Burdwan for his support, advice and helpful suggestions during compilation of this manuscript.
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