Locomotory modes in the larva and pupa of Chironomus plumosus (Diptera, Chironomidae)

The locomotory kinematics of Chironomus plumosus larvae and pupae were investigated in order to determine how different locomotory techniques may be related to (a) possible underlying patterns of muscle activation and (b) the particular lifestyles and behaviours of these juvenile stages. Larvae display three independent modes of motile activity: swimming, crawling and whole-body respiratory undulation. Swimming and respiratory undulation involve the use of metachronal waves of body bending which travel in a head-to-tail direction. Whereas swimming is produced by side-to-side flexures of the whole body, respiratory undulation employs a sinusoidal wave. Crawling appears to result from an independent programme of muscle activation. Instead of a longitudinally transmitting metachronal wave of body flexure, a simultaneous arching of the body, combined with the alternating use of the abdominal and prothoracic pseudopods as anchorage points, produces a form of locomotion analogous to caterpillar-looping. Larval swimming has a set speed and rhythm and is an 'all-or-nothing' locomotory manoeuvre, but the neural programme controlling larval crawling is adaptable; switching from a less to a more slippery substrate resulted in a shorter, faster stepping pattern. The pupa displays two swimming modes, somersaulting and eel-like whole-body undulation, the former being principally a brief, escape manoeuvre, the latter being a faster form of locomotion employed to deliver the pupa to the surface prior to adult emergence. Comparison with the pupa of the culicid Culex pipiens shows that this insect also uses the somersault mechanism but at a higher cycle frequency which produces a faster swimming speed. This appears to be related to differences in lifestyle; the surface-living culicid pupa is exposed to greater predator threat than the bottom-dwelling chironomid pupa, and consequently needs a faster escape.     

Divergence of karyofunds in sibling species of the plumosus group (Diptera: Chironomidae)

Cytogenetic differentiation of eight sibling species of the plumosus group was examined. The karyofunds of these sibling species were shown to diverge incompletely. In each species karyofund, the banding sequences homologous to those of the remaining species of this group were revealed. The number of banding sequences that displayed interspecific homology varied from 3 to 13 per species karyofund. In a species karyotype, the homologous sequences were localized to chromosome arms 1-6. Both similar and contrasting frequencies of homologous banding sequences were observed in karyofunds of different sibling species. The average cytogenetic distance between sibling species of the plumosus group was 2.618 +/- 0.400. The presence of species-specific banding sequences, the absence of homologous banding sequences in some chromosome arms of the karyotype, and different frequencies of the homologous banding sequences determined the cytogenetic divergence of the sibling species.     

Cytogenetic studies of Chironomus circumdatus from India (Diptera: Chironomidae)

Chironomus circumdatus is one of the most common and wide-spread species in India. Cytogenetic studies pertaining to the mitotic and polytene chromosomes, nucleolar organizer regions, C-banding and naturally occurring chromosomal polymorphism have been carried out for the first time in this Indian species. Altogether seven inversions comprising six paracentric and one pericentric are detected in the Indian populations. The distribution of inversions in relation to different environmental conditions is discussed.     

Cytogenetic structure of Chironomus plumosus (L.) (Diptera, Chironomidae) population in regard to the trophic state of the Curonian Lagoon, Baltic Sea

It is shown that the cytogenetic structure of the Chironomus plumosus (L.) population in the Curonian Lagoon has not changed significantly over the last three decades. The existing cytogenetic structure supported not only population survival, but also its proliferation in new environmental conditions caused by an increase of the trophic status of the water body. The predominance of homozygotes pluB2.2 was marked in karyofund. Both the oxygen deficiency periodically occurring in the Curonian Lagoon and the fact that the population dwells at the periphery of the species areal in the Palearctic could be reasons for this phenomenon.     

C-banding in the polytene chromosomes of species of the plumosus group (Diptera,Chironomidae) and their experimental hybrids

The localization and amount of heterochromatin in the plumosus group were studied, including the species Chironomus plumosus L., C. vancouveri and C. balatonicus. The appearance of C bands of Chironomus plumosus in several European populations is traced. The role of the C heterochromatin in the differentiation of this species is discussed. From the evolutionary point of view the Swiss populations, in which large centromere heterochromatin blocks have been discovered, are more varied as to the amount of heterochromatin. The importance of duplications for this process is pointed out. The chromosomes of the individuals from C. vancouveri and C. balatonicus have centromeric, telomeric and interstitial heterochromatin. The centromeric heterochromatin is represented by thin C-bands. The particularities in the appearance of C heterochromatin in C. vancouveri and C. balatonicus reflect the structural peculiarities of their chromosomes. The change in the euchromatin regions in these forms is discussed in the light of transformation of euchromatin to heterochromatin in the process of evolution.The appearance of heterochromatin in hybrids (between populations and between species) created experimentally is traced. A change has been discovered in the appearance of heterochromatin in the hybrids compared to the initial parent forms. This difference is expressed more strongly in inter-species hybrids than in interpopulation hybrids of C. plumosus.     

Peculiarities of structural organization of hemoglobin of Chironomus plumosus L. (Diptera: Chironomidae)

In Ch. plumosus and Ch. riparius there were revealed various structural hemoglobin variants including monomers, dimers, trimers, tetramers, hexamers, and octamers. The multitude of the protein organization ways seems to be based on diversity of the monomers with the approximately equal molecular weight from 12 to 16 kDa in Ch. plumosus and from 11.8 to 15.2 kDa in C. riparus. In PAGE with 8 M urea, the hemoglobins were aggregated into high molecular complexes with mol. weights about 260 and 180 kDa in Ch. plumosus and about 523 and 174 kDa in C. riparus.     

Seasonal variability of the karyotype structure of Chironomus plumosus (Diptera, Chironomidae) from a biotope of Kaliningrad

Chironomus plumosus larvae from the polluted Shkolnoe lake, Kaliningrad, have 2n = 8 and 2n = 8 + B. In winter season we found 11 types of hetero- and homozygous inversions in A, B, C, D, E, and F arms whereas in summer season we registered 7 types of the same inversions in A, B, C, D, and E arms. All inversions with exception of the inversion in arm C correspond to Hardy-Weinberg equilibrium. The arm IVG shows homozygous increase of centromeric heterochromatin more frequently in summer than in winter (34.4% as compared with 1.8%). The arm E has asynapsis 2 times less frequently in summer than in winter (21.4% as compared with 44.6%).     

Intercontinental karyotypic differentiation of Chironomus entis Shobanov, a Holarctic member of the C. plumosus group (Diptera, Chironomidae).

Analysis of banding sequences of polytene chromosomes in Palearctic (Russian) and Nearctic (North American) Chironomus entis shows strong karyotype divergence between populations on the two continents. Four out of seven chromosomal arms in the North American C. entis karyotype are characterized by sequences found only in the Nearctic. In total, 44 banding sequences are now known for this species across the Holarctic, including 22 exclusively Palearctic, 6 Holarctic, and 16 exclusively Nearctic sequences. The degree of cytogenetic differentiation between Palearctic and Nearctic C. entis populations is an order of magnitude greater than differentiation among populations within either continent, but is only one third as great as the cytogenetic distance between the sibling species C. entis and C. plumosus. C. entis is the only sibling species of C. plumosus uncovered during cytological identification of Chironomus species from more than 50 North American lakes, indicating that the plumosus sibling-species group is much smaller in the Nearctic than in the Palearctic, where a dozen sibling species are known. Cytogenetic distance values calculated between Nearctic and Palearctic representatives of both C. entis and its sibling species C. plumosus are similar, but result from different patterns of karyotype divergence. New World C. entis is distinguished from Old World populations by the 16 uniquely Nearctic sequences, four of which occur in the homozygous state. In contrast, North American C. plumosus has fewer uniquely Nearctic sequences, and only one that occurs as a homozygote. However, four chromosomal arms in C. plumosus that are polymorphic in the Palearctic show fixation, or near fixation, of Holarctic sequences in the Nearctic C. plumosus karyotype. Thus, both the fixation of Holarctic sequences, and the occurrence or fixation of distinctly Nearctic sequences, contribute significantly to karyotype divergence. Patterns of karyotype divergence in Palearctic and Nearctic populations of different Holarctic chironomid species are discussed relative to intercontinental cytogenetic differentiation in other dipterans.     

A phylogenetic study of sex determiner location in a group of Australasian Chironomus species (Diptera,Chironomidae)

Sex determination in a group of phylogenetically related Chironomus species, of the pseudothummi complex, from south-eastern Australia and New Zealand is male heterogametic, controlled by a male determiner. The male determiner has been located at least to the level of the chromosome arm in most members of this phylogenetic group. It varies in location among many of the species and there are some phylogenetic patterns discernable, which are discussed in relation to the possible origin of the sex determiner. There is a group of species, Ch. oppositus ff. oppositus and whitei, Ch. australis, Ch. alternans a and Ch. alternans c, which appear to be central to this phylogeny, in which the sex determiner is located near the centromere of the CD chromosome, the most common location in the Australasian group. This is different from the most common location, arm F, of the thummi complex in Europe and North America. There is also a group, comprising Ch. oppositus f. tyleri, Ch. cloacalis, Ch. alternans b and Ch. nepeanensis, in which the sex determiner is on arm G. The arm A sex determiners, found in Ch. tepperi, Ch. oppositus ff. whitei and connori, and Ch. occidentalis, may be of common origin or they may be independently derived, as must be the arm B (Ch. duplex) and arm F (Ch. oppositus f. whitei) sex determiners. In Ch. oppositus f. whitei, four different chromosomal locations for the sex determiner have been identified. It is not yet clear whether these represent an unstable polymorphism or indicate the existence of cryptic subgroupings within this form. Although the location of the sex determiners can be assigned to particular chromosome arms, the precise location cannot be determined, therefore the assumption of common origin may not always be correct. Also, this uncertainty means that it is impossible at present to differentiate between a complex system of sex determination and the possibility of a translocatable sex determiner as explanations of the variability in sex determiner location. The forms of Ch. oppositus are redefined and renamed to avoid confusion caused by the previous names.It is a pleasure to dedicate this paper to Professor Hans Bauer, a pioneer of chironomid karyosystematics, on the occasion of his 80th birthday     

Karyofund of Chironomus plumosus (Diptera, Chronomidae) in the Palearctic region

Quantitative and qualitative analyses of chromosomal polymorphism in 38 Palearctic populations of Chironomus plumosus were made. It was shown that most of the populations studied were highly polymorphic: in average 63.2 +/- 4.0% of larvae were heterozygous for inversions with 0.95 + 0.08 heterozygous inversion per larvae. Polymorphism on the size of centromeric heterochromatin and the presence of B-chromosomes were observed in many populations studied. The karyofund of Ch. plumosus in Palearctic was estimated. In total 35 banding sequences were found in Palearctic Ch. plumosus. Fifteen banding seguences have been described for the first time. On mapping the used banding sequences, we employed the conventions of Keyl (Keyl, 1962; Devai et al., 1989) and Maximova (Maximova, 1976; Shobanov, 1994a) for arms A, C, D, E and F, and the conventions of Maximova for arms B and G.     

Chromosomal and genomic mutations in Chironomus plumosus (L.) (Diptera, Chironomidae) from Novozybkov raion of Bryansk oblast

The data on karyopool analysis of Chironomus plumosus from Novozybkov raion of Bryansk oblast, radioactively contaminated as a result of the Chernobyl accident, are presented. In the karyopool of this population, four types of heterozygous inversions (A1.2, B1.2, C1.2, D1.2) and a structurally small rearrangement in arm D, which is thought to be also associated with inversion, were identified. For inversions A1.2, C1.2, and structurally small rearrangement in arm D the cases of somatic mosaicism were described. The mean number of inversions per individual constituted 0.78. The number of genotypic combinations was 13. In 5% of the individuals chromosome B was identified, while 15.6% were polyploid (3n). In 1.8% of triploids somatic mosaicism for the level of polytene chromosomes was observed. Most of the larvae were characterized by partial asynapsis of the homologs in different regions of chromosome III. Specific features of the Chironomus plumosus karyopool from Novozybkov are thought to be associated with the habitation of a number of generations of this population on radioactively polluted territory.     

Karyofunds of Chironomus plumosus (L.) (Diptera, Chironomidae). V. terminal and interstitial populations

The study of polytene chromosomes of Chironomus plumosus from Rybinsk reservoir, its inflows and surrounding water bodies allow to mark out two type's of population. Terminal populations--presented by karyoforms with concrete limited composition of genotype combinations and zygotic arm combinations, which adapted to specific complex of the habitat factors. The immediate exchange of genes between different terminal populations is absent (even they placed in nearness) from allochronie of life cycles and impossibility of larvae-immigrants to finish ontogenesis in unfit for they habitat conditions. Interstitial populations inhabit the biotopes, where the karyoforms of two or more terminal populations can to finish ontogenesis and to reproduce. The genetic exchange between terminal population implement through interstitial populations.     

Resistance to anoxia of Chironomus plumosus and Chironomus anthracinus (Diptera) larvae

Survival of the midge larvae Chironomus plumosus and C. anthracinus in anoxia at 4°C was investigated. C. plumosus survived about twice as long as C. anthracinus . The corresponding LT 50 values were ca 205 and 100 d. There was no statistically significant difference between the survival in anoxia and in aerated water, which indicates that the main reason for death in anoxia is not the absence of oxygen. This main reason is presumed to be starvation. The presence of undissociated H₂S in low concentrations (ca 1.2 mg 1⁻¹) did not influence the survival. In anoxia the larvae were usually motionless and did not feed but they increased in weight due to uptake of water. During 43 d C. plumosus and C. anthracinus increased ca. 10%.     

A review of the genus Chironomus (Diptera,Chironomidae)

Analysis of the banding pattern of the salivary gland chromosomes of Chironomus tepperi indicates that, despite a somewhat modified male hypopygium, the relationships of this species are close to the other Australian species of the genus, particularly to Ch. oppositus. No inversion polymorphism has been found in Ch. tepperi and this, together with the relatively high chiasma frequency as measured at metaphase I, would appear to be an adaptation to provide genetic variability necessary for its colonizing ability.     

Karyotype of the midge Chironomus heterodentatus Konstantinov from the group Obtusidens (Diptera, Chironomidae)

The karyotype of Chironomus heterodentatus, belonging to the obtusidens-group described by Konstantinov from the Volga in 1956, was studied in detail. Combinations of chromosomal arms are AB, CD, EF and G (cytocomplex thummi). The Ch. heterodentatus karyotype cleary differs from those of other members of the obtusidens-group with a species-specific banding pattern in arms A, B, C and D. Chromosomal polymorphism on homo- and heterozygous inversions was found in arms, A, B, D, E and G. 19 inversion banding sequences and their 27 genotypic combinations have been recorded. The shortest arm G is highly polymorphic. Heterozygotes on the Balbiani ring activity were found in arm G along with homo- and heterozygotes on inversions. B-chromosomes with a frequency equal to 2.7-25.0% were recorded in some Volga populations.     

Cytogenetic characteristics of a population of Chironomus riparius Meigen 1804 (Diptera, Chironomidae) from a polluted Po river station

A population of Chironomus riparius from a Po river station near Moncalieri (a trace-metal polluted station) was studied. In this population was established a great variability of band structure of polytene chromosomes as well as paracentric heterozygous inversions, deletions, deficiencies, partial breaks, diploid chromosome fragments, and changes in functional activity and appearance of heterochromatin. In arms A through F, some bands had an increased size compared to the standard chromosomic map. Some bands appeared in a heterozygous or normal homozygous state or were amplified. In all arms, many condensed stable bands appeared in the decondensed state when compared to the standard map. Asynaptic zones in arms E and G as well as heterozygous Balbiani rings and NORs were established. Very often the 4th chromosome was almost completely heteropycnotic and looded like a pompon chromosome. For the first time in this species, a high frequency of ectopic pairings of different arms was observed. Telomeric regions involved in ectopic pairings had a granular appearance, as did some centromeres. The hypothesis is advanced that such a high frequency of structural rearrangements could be correlated with genomic distribution of specific mobile elements.     

DNA barcoding supports reclassification of Japanese Chironomus species (Diptera: Chironomidae)

Non‐biting midges (Diptera: Chironomidae) adapt to species‐specific environmental conditions and hence are promising bioindicators for aquatic and ecotoxicological monitoring. Although their utility for these purposes was historically limited by difficulties in their morphological identification, DNA barcoding offers a possible solution. Here, eight Japanese species of the genus Chironomus, which is characterized by its worldwide distribution and abundance among Chironomidae, were subjected to DNA barcoding using cytochromec oxidase subunit I (COI). To examine whether this DNA barcode is a useful indicator for Japanese species of Chironomus, we calculated genetic distances within and between the COI sequences of Chironomus species both from this study and worldwide and constructed phylogenetic trees. Based on 415 bp COI sequences and the Kimura two‐parameter model, the average genetic distances within 37 species and between 72 species were 2.6% and 17.2%, respectively. Although the ranges of genetic distances within and between species overlapped from 0.8% to 17.3%, 99.7% of average genetic distances between species were >3.0%. Some of this overlap is attributable to distances within species that were “too large” as well as those between species that were “too small”. Of eight Japanese species examined, two showed genetic distances between species that were below a 3.0% threshold, and four had distances within species that were greater than 3.0%. These results suggest a possible reclassification of these species and the need for further sampling to unveil biogeographic variations among different countries and regions.     

Seasonal Dynamics of a Chironomus plumosus (L.) (Diptera,Chironomidae) Population from a Fish Pond in Southern Bohemia

A population of Chironomus plumosus (L.) from a fish pond has been investigated for two years. Quantitative benthic samples and emergence traps were used to study the seasonal dynamics. Effect of some factors on the duration of development was studied in laboratory experiments. Data on larval abundance, length frequency analysis and emergence of adults suggest a bivoltine life cycle. The population of C. plumosus investigated showed a pronounced tendency to dormancy, even under long-day conditions. Factors regulating the bivoltine life cycle are discussed.