Cytogenetic Features of Blackfly Wilhelmia paraequina Puri (Diptera: Simuliidae) from Armenia

Three blackfly Wilhelmia paraequina populations of Armenia (rivers Debet and Megriget, channel Megri) were studied. 2n = 6: IS + IIL, IIS + IL, IIIS + IIIL. Cytological maps of polytene chromosomes were constructed. High inversion polymorphism (95.63%) was observed, with 2.6 inversions per individual. Three inversions proved to be associated with male development: Y₁ was characterized by a combination of two heterozygous inversions, IIL-3 + IIL-1, while Y₂ had another combination of heterozygous inversions, IIL-5 + IIL-3. The X chromosome had a standard homozygous IIL band pattern. The populations were shown to be similar in autosomal polymorphism. A tendency for differentiation was observed with respect to frequencies and types of sex-linked inversions: the sex determination system was Y₁X–XX in the Debet and Megriget populations and Y₂X–XX in the channel (Megri) population. On the strength of these findings, one W. paraequina morphotype was assumed to involve two cytotypes, A (Debet and Megriget populations) and B (Megri population).     

Cytogenetic analysis of the noxious bloodsucker Boophthora erythrocephala (Diptera: Simuliidae) from different geographic zones

Boophthora erythrocephala is a widely spread species inhabiting both small polluted water-bodies and large rivers. Three populations from Samara region in Russia and from Donetsk and Chernigov regions in the Ukraine have been studied. Five populations of this species in Russia were described earlier (Polyanskaya, Tsapygina, 1968; Petrukhina, 1972). For identification of this species the chromosome map of Adler and Werner (personal communication) was used. Comparison of 8 populations studied was performed. All of them belong to the species B. erythrocephala but everyone has its own individual characteristics. Inversion polymorphism was observed in each population comprising from 88 to 100 %. Mean number of heterozygous inversions per individual is low fluctuating around 3.6. There is one inversion in IIS observed in all populations that proves panmixy among them. Other inversions occur in a part of the individuals.     

Cytogenetics of Simulium siamense Takaoka and Suzuki, 1984 (Diptera: Simuliidae) in northeastern Thailand

We analysed salivary gland polytene chromosomes of 796 larvae from 17 populations of Simulium siamense in northeastern Thailand. Seventeen floating and two fixed chromosome inversions were recorded. Three cytoforms (A, F and G) were recognised and two of them are new (F and G). Cytoform F is distinguished by a fixed inversion on the long arm of chromosome II (IIL-8) and cytoform G by fixed inversions on the long arm of chromosome II (IIL-8) and short arm of chromosome III (IIIS-2). Significant departures from Hardy–Weinberg equilibrium due to heterozygote deficiency in geographically intermediate populations and absence of shared polymorphic inversions of the cytoforms indicate separation of the gene pool. Morphometric analysis of the larvae revealed significant differences in body length (F = 5.00, p =0.007) and head capsule width (F = 4.68, p = 0.010) among cytoforms.     

Study of the karyotype of the malaria mosquito]

A study of the karyotype composition of 22 geographycally removed natural populations of the malarial mosquito Anopheles messeae Fall revealed a chromosomal polymorphism characteristic of the whole species. 13 paracentric inversions were detected, no other gene arrangements being discovered. All the inversions can be divided into two classes: 1) wide spread inversions in hetero- and homozygous forms; 2) endemic ones discovered in individual populations in the form of heterozygotes. According to the presented photomap of salivary gland chromosomes, the inversion localization are as follows: IL1 (2a--4a), IL2 (1c--4a), IL3 (2a--3b), IL4 (1c--3b), IIR1 (7c--12c), IIR2 (10b--14b), IIL1 (15b--17a), IIIR1 (24a--26c), IIIR2 (23a--24c), IIIR3 (23b--25c), IIIR4 (27c--29c), IIIR5 (27d--30c), IIIL1 (34a--39d).     

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.     

Speciation and chromosomal evolution of the Baikalian endemic chironomids of the genus Sergentia Kief. (Diptera, Chironomidae): Karyotype divergence and chromosomal polymorphism in the populations of deep-water species Sergentia nebulosa Linevitsh et al. and Sergentia assimilis Proviz V. et Proviz L.

Specific karyotype structure and chromosomal polymorphism was investigated in the populations of Sergentia nebulosa Linevitsh et al., 1984 and Sergentia assimilis Proviz V. et Proviz L., 1999, the deep-water endemic chironomid species (Diptera, Chironomidae) from the Baikal Lake. The distinguishing feature of the karyotypes of these species, compared to the other Baikalian Sergentia, is well-developed nucleolus in region 6 of arm C. Both species display the presence of interspecific population polymorphism, determined by the structure of this arm. In some populations, chromosome regions from 4 to 6 contain a homozygous inversion, which is absent in the other populations. The distinguishing karyotype feature of S. assimilis, which shares fluctuating homozygous inversions with the other species, is the presence of two species-specific homozygous inversions. These are the secondary overlapping inversion in arm A, regions 2 to 7, and the inversion in regions 4 to 10 of arm G. Both species of interest contain nucleolus organizer in region 10 of arm G. In populations of S. nebulosa, six heterozygous inversions localized in arms A, B, C, F, and G were discovered. The highest number of heterozygotes for inversions (71%) was observed in the population from Southern Baikal. In arm B of S. assimilis, one heterozygous inversion and heterozygosity for nucleolus organizer in the chromosome region 16 was detected. Chromosomal evolution of Baikalian Sergentia, and the role of inversion polymorphism in the population adaptation is discussed. Original Russian Text ? V.I. Proviz, 2008, published in Genetika, 2008, Vol. 44, No. 12, pp. 1627–1637.     

Cytogenetics of natural populations of Drosophila

D. bipectinata and D. malerkotliana differ from each other by three overlapping inversions in IIL, two included inversions in IIIL and two overlapping inversions in IIIR. These inversions were analysed on the basis of the salivary chromosome maps of D. malerkotliana. Bock's (1971) data revealed that the four members of the bipectinata species complex differ from each other with respect to overlapping inversions. The reason why the ancestral population which may have been heterozygous for common inversions split into at least four groups, each leading to the formation of a new species, and the possible mechanism of the origin of sexual isolation between the groups is discussed.     

Speciation and chromosomal evolution of the Baikalian endemic chironomids of the genus Sergentia Kief. (Diptera, Chironomidae): karyotype divergence and chromosomal polymorphism in the populations of eurybathic species Sergentia flavodentata Tshern. and littoral species Sergentia baicalensis Tshern

Specific karyotype structure and chromosomal polymorphism was investigated in the populations of the Baikalian endemic chironomid species, eurybathic Sergentia flavodentata Tshernovskij, 1949 and littoral Sergentia baicalensis Tshernovskij, 1949. In addition to two fluctuating homozygous inversions in arms A and E, both species were characterized by species-specific karyotype features, namely, nucleolar organizer in the region 1 of chromosome IV in S. flavodentata, and inversion in arm F in S. baicalensis. Moreover, S. baicalensis demonstrated the presence of intraspecific population polymorphism. The populations differing in the presence of secondary overlapping inversion in arm A were found. The highest number of chromosomal rearrangements (7) was detected in S. flavodentata. Most of these rearrangements (six) were found in the population from the underwater thermal spring at a depth of 420 to 430 m (Northern Baikal, Frolikha Bay). In the populations from Middle and Southern Baikal, a rare pericentric inversion in chromosome I was described. In S. baicalensis, in addition to two common heterozygous inversions in arms C and F, disturbance of homologous pairing in different regions of the remaining arms were often detected. Stable chromosomal polymorphism preserved during 13 years in the population of S. flavodentata from the region of hydrothermal venting, serves as an evidence of high genetic plasticity of the species, which favors successful colonization of different Baikal depths and biotopes.     

Inversion polymorphism in the population of Camptochironomus tentans from Kaliningrad city

This paper present the map of polytene chromosomes and inversion polymorphism of widely distributed Chironomidae species, Camptochironomus tentans, from the most western locality of Russia--Kaliningrad city. Chromosomes banding pattern is designated according to Beermann, 1955. Only one larva (2.9%) had a standard banding pattern, and karyotypes of the rest (97.1%) were polymorphic. We have found 2.0 heterozygous inversions per individual, and the frequency of hetero- and homozygous inversions, taken together, amounted to 2.2 per individual. 17 inversion banding patterns and 20 genotypic combinations of these patterns were found. It was shown that the most frequent inversions in this population were identical to these in European populations.     

Chromosomal polymorphism of the unique Baikal deep-water population of Sergentia flavodentata (Diptera, Chironomidae) from the region of hydrothermal venting

The subject of this study is chromosomal polymorphism of a population of Baikal endemic species Sergentia flavodentata (Diptera, Chironomidae) from the Frolikh Bay region of hydrothermal venting at a depth of 420-430 m. In 35-67% of larvae, six heterozygous inversions were found to be preserved for 13 years (materials of 1991, 1994, 1996, 1999, 2003). The number of zygotic combinations reached 14; 0.5-0.7 inversions per individual was observed. It is suggested that stable chromosomal polymorphism characteristic of this species is nonrandom, being a mechanism of the population adaptation to the specific conditions of hydrothermal venting community based on bacterial biochemosynthesis.     

Seasonal changes of chromosomal inversion polymorphism in a Drosophila subobscura natural population from southeastern European continental refugium of the last glaciation period

Chromosomal inversion polymorphism was studied in a natural population of Drosophila subobscura from Djerdap gorge of Danube river, that represents one of southeastern European continental refugiums during last glaciation period. The existence for changes of chromosomal inversion polymorphism during different periods of time was tested. Observed very high abundance of chromosomal arrangements and lower IFR value grouped this population to central European populations. It was characterized with O7 and O(3+4+5) arrangements, not found in other populations from the region analyzed. Except in this population from warmer habitat, observed E(1+2+9+12) chromosomal arrangement was found, in the region analyzed, only in one population from Mediterranean refugial area. Many similarities have been observed between population from Djerdap gorge and other populations from the region analyzed, indicating that process of post-glacial migration may have started from this southeastern European refugium area. Seasonal changes in the frequencies for Est and O(3+4+7) chromosomal arrangements where Est was more frequent during the spring period and O(3+4+7) during the summer period were found. The changes for J-chromosome (Jst/Jst), E-chromosome (Est/E(1+2+9); E(8)/E(1+2+9) and O-chromosome (Ost/O(3+4+22); O(3+4)/O(3+4); O(3+4+1)/O(3+4+7) whose karyotypic combination frequencies were associated with seasonal changes were also found. These changes most probably represent associations with seasonal climatic changes in the population analyzed, supporting adaptivity of chromosomal inversions in D. subobscura.     

The Fertility Effects of Pericentric Inversions in Drosophila Melanogaster

Heterozygotes for pericentric inversions are expected to be semisterile because recombination in the inverted region produces aneuploid gametes. Newly arising pericentric inversions should therefore be quickly eliminated from populations by natural selection. The occasional polymorphism for such inversions and their fixation among closely related species have supported the idea that genetic drift in very small populations can overcome natural selection in the wild. We studied the effect of 7 second-chromosome and 30 third-chromosome pericentric inversions on the fertility of heterokaryotypic Drosophila melanogaster females. Surprisingly, fertility was not significantly reduced in many cases, even when the inversion was quite large. This lack of underdominance is almost certainly due to suppressed recombination in inversion heterozygotes, a phenomenon previously observed in Drosophila. In the large sample of third-chromosome inversions, the degree of underdominance depends far more on the position of breakpoints than on the inversion's length. Analysis of these positions shows that this chromosome has a pair of ``sensitive sites' near cytological divisions 68 and 92: these sites appear to reduce recombination in a heterozygous inversion whose breakpoints are nearby. There may also be ``sensitive sites' near divisions 31 and 49 on the second chromosome. Such sites may be important in initiating synapsis. Because many pericentric inversions do not reduce the fertility of heterozyotes, we conclude that the observed fixation or polymorphism of such rearrangements in nature does not imply genetic drift in very small populations.     

Temephos-resistant larvae of Simulium sanctipauli associated with a distinctive new chromosome inversion in untreated rivers of south-western Ghana

Larvae of the Simulium damnosum Theobald complex (Diptera: Simuliidae) were sampled in June 1996 from two sites in south-west Ghana where larviciding has not been applied: Sutri Rapids on the Tano river (05 degrees 23 minutes N 02 degrees 38 minures W) and Sekyere-Heman on the Pra river (05 degrees 11 minutes N 01 degrees 35 minutes W). All specimens were identified as Simulium sanctipauli Vajime & Dunbar sensu stricto (Diptera: Simuliidae). Bioassays with temephos (organophosphorus larvicide employed by the Onchocerciasis Programme for systematic treatment of most rivers across West Africa since the 1970s) showed about five-fold resistance in the Tano population (LC95 2.37-3.14 mg/L) and slight tolerance to temephos in the Pra population (LC95 0.67-0.76 mg/L), vs. the diagnostic concentration of 0.625 mg/L. Larval salivary polytene chromosomes of S. sanctipauli showed fixed inversions 1S-24/24, standard IIL-6 and a new inversion IL/36 polymorphism at Sutri on the Tano. These karyotype characteristics differ from those of temephos-resistant S. sanctipauli in rivers of C te d'Ivoire and other sites on the Tano in Ghana. Thus, temephos resistance in S. sanctipauli at Sutri is associated with distinct chromosomal configurations, showing that immigration was unlikely. This resistance could have been locally selected by exposure of S. sanctipauli larval populations to agrochemicals run-off from cocoa, coffee and oil plantations flanking the rivers.     

Comparative cytogenetic variability of polytene chromosomes from salivary glands of Chironomus riparius Mg., 1804 (Diptera, Chironomidae) from two polluted biotopes in Bulgaria and Russia

The structural-functional variation of Chironomus riparius salivary gland polytene chromosomes was studied in two geographically isolated Palearctic regions, Bulgaria (village Pancharevo) and Russia (St. Petersburg). The two biotopes, where larvae were collected, were polluted with various heavy metals from anthropogenic sources. Hereditary paracentric heterozygous inversions were characteristic of the Russian population, whereas somatic paracentric or pericentric heterozygous inversions were more common in the Bulgarian one. All inversions occurred at low frequencies. Other aberrations found in the two populations included somatic deletions resulting in a pompon structure of chromosome IVG, heterozygous translocation between chromosomes IVG and IIIF, enlargement of individual disks, and the appearance of a heterozygous block close to the centromere of chromosome IVG. In addition, changes in functional activity of the nucleolus organizer and Balbiani rings (BRc/BRb) were observed. Several aberration breakpoints proved to coincide with satellites of the Alu and Hinf families.     

Chromosomal inversion polymorphism of Anopheles funestus from forest villages of South Cameroon.

The polymorphism of paracentric inversions of Anopheles funestus polytene chromosomes was studied in three villages (Nkoteng, Obala, and Simbock) located in a forest area of South Cameroon in order to analyse the genetic structure of these populations. A total of 146-210 chromatids could be scored from specimens collected over about two years. A low degree of chromosomal polymorphism was observed with two floating inversions on chromosomal arm 2 (2h and 2d), and three fixed arrangements on arms 3 (3a and 3b), and 5 (5a). Such arrangement of inversions has never been recorded elsewhere so far. The chromosome analysis indicated that the population from Obala was in Hardy-Weinberg equilibrium, whereas the samples from Nkoteng and Simbock showed a significant excess and deficit of heterokaryotypes, respectively. Significant differences in inversion frequencies on chromosomal arm 2 among villages lying in contrasting eco-climatic settings suggested an adaptive role of these inversions.     

Speciation and chromosomal evolution of the palaearctic species of chironomids of the genus Sergentia Kieffer (Diptera, Chironomidae): Chromosome polymorphism in the populations of S. baueri Wülker et al., 1999, S. prima Proviz et al., 1997, and S. electa Proviz et al., 1999 from the Irkutsk reservoir

Chromosome polymorphism was studied in populations of three Palaearctic species of chironomids of the genus Sergentia Kieffer (Diptera, Chironomidae) from the Irkutsk reservoir: S. baueri Wülker et al., 1999, S. prima Proviz et al., 1997, and S. electa Proviz et al., 1999. It was found that in S. baueri heterozygous individuals constitute 72% of the population with 0.8 of inversion per individual. In total, three inversion sequences were detected. The most widespread was the inversion in regions 7–12 of the arm IIIR with the borders similar to those of the inversion in the Far East S. baueri populations and in populations of other species of the genus. In S. prima, heterozygotes constituted 60%, with the number of inversions per individual was 1.3. Seven inversion sequences were revealed, of which three were found in chromosome IV. In the S. electa population, only one inversion sequence was observed in chromosome III in 36% of larvae (0.4 of inversion per individual). The level of chromosome polymorphism in the populations of the Palaearctic species is comparable with that in the populations of endemic Sergentia from the Baikal Lake. Ten rearrangements were revealed in each of the two groups, but similar borders of chromosome regions were established only for two inversions. In the studied Sergentia species, chromosome IV has the most variable structure.     

Cytogenetics of the Anopheles gambiae complex in Sudan, with special reference to An. arabiensis: relationships with East and West African populations

The species composition of malaria vector mosquitoes belonging to the Anopheles gambiae complex (Diptera: Culicidae) from >40 localities in Sudan, representing most ecological situations, was determined by analysis of ovarian polytene chromosomes. Of 2162 females, 93% were identified as An. arabiensis Patton and 7% were An. gambiae Giles sensu stricto. No hybrids were found between the two species. Anopheles arabiensis occurred in all but two sites, whereas An. gambiae s.s. was effectively limited to the southernmost, more humid localities. For chromosomal paracentric inversions, the degree of polymorphism was low in An. gambiae s.s. (inversions 2La, 2Rb and 2Rd), higher in An. arabiensis (inversions Xe, 2Ra, b, bc, d1, s; 3Ra, d). Anopheles gambiae samples from Sudan were all apparently panmictic, i.e. they did not show restricted gene flow such as observed among West African populations (interpreted as incipient speciation). Chromosomal inversion patterns of An. gambiae in southern Sudan showed characteristics of intergrading Savanna/Forest populations similar to those observed in comparable eco-climatic situations of West Africa. Anopheles arabiensis was polymorphic for inversion systems recorded in West Africa (2Ra, 2Rb, 2Rdl, 3Ra) and for a novel 2Rs polymorphism, overlapping with inversion systems 2Rb and 2Rd1. Samples carrying the 2Rs inversion were mostly from Khashm-el-Girba area in central-eastern Sudan. In the great majority of the samples all polymorphic inversions were found to be in Hardy-Weinberg equilibrium. Sudan populations of An. arabiensis should therefore be considered as generally panmictic. Anopheles arabiensis shows more inversion polymorphism in west than in east African populations. Sudan populations have more evident similarities with those from westwards than those from eastwards of the Great Rift Valley. The possible influence of the Rift on evolution of An. arabiensis is discussed.     

The geographic varyability of the polytene chromosome banding sequence of non-biting midje Chironomus pseudothummi str. (Diptera, Chironomidae)

The karyotypes and chromosomal polymorphism of Chironomus pseudothummi were investigated in different parts of its range. It was established that chromosomal variability in the natural populations of this species was represented mainly by the inversion polymorphism of arm G. Only rare and unique inversions were found as heterozygous in arms C, D, and E. In total, the 14 banding sequences of polytene chromosomes form the banding sequences pool of Ch. pseudothummi. Geographic differences in distribution of chromosomal banding sequences throughout the range were established. The presence of banding sequences pstG1 and pstG2 is characteristic of European populations. Banding sequence pstG1 was completely vanished with simultaneous increase in frequency of pstG2 and appearance of new inversion banding sequence pstG3 in Siberian populations. The differences in the set of the rare and unique inversions in arms C, D, and E between west-European and west-Siberian populations were revealed.     

UDP-glucuronosyltransferase 1A4 (UGT1A4) polymorphisms in a Jordanian population

Glucuronidation is one of the most important phase II metabolic pathways. It is catalyzed by a family of UDP-glucuronosyltransferase enzymes (UGTs). One of the subfamilies is UGT1A. Allele frequencies in UGT1A4 differ among ethnic groups. The aim of this study was to determine the allelic frequency of two most common defective alleles: UGT1A4*2 and UGT1A4*3 in a Jordanian population. A total of 216 healthy Jordanian Volunteers (165 males and 51 females) were included in this study. Genotyping for UGT1A4*1, UGT1A4*2 and UGT1A4*3 was done using a well established polymerase chain reaction-restriction fragment length polymorphism test. Among 216 random individuals studied for UGT1A4*2 mutation there were 26 individuals who were heterozygous, giving a prevalence of 12% and an allele frequency of 6.5%. Only one individual was homozygous for UGT1A4*2. The UGT1A4*3 mutation was detected as heterozygous in 9 of 216 individuals indicating a prevalence of 4.2% and allele frequency of 3.5%. Three individuals were homozygous for the UGT1A4*3 indicating a prevalence of 1.4%. The prevalence of UGT1A4*2 is similar to the Caucasians but different from other populations whilst the UGT1A4*3 prevalence in the Jordanian population is distinct from other populations. Our results provide useful information for the Jordanian population and for future genotyping of Arab populations in general.     

Chromosome Studies in Wild Populations of D. MELANOGASTER

Chromosome studies of wild D. melanogaster populations from Missouri, Mississippi, Louisiana and Texas uncovered 58 inversions. Six were common and cosmopolitan; 52 were new, rare and generally endemic. In one of two Missouri populations tested, structurally heterozygous females carried significantly more sperm at capture than did the homozygotes. In both populations comparisons of wild sperms with the females carrying them indicated significant positive assortative mating and an excess production of homozygotes among the F₁ progeny. Wild females structurally heterozygous in up to three major autosomal arms showed no associated nondisjunctional egg lethality; those heterozygous in all four arms produced from 0% to 24% dead eggs, suggesting the presence of intrapopulational gene modifiers of meiosis. Texas populations supported on windfall citrus fruit showed a slight but significant difference in inversion frequencies between flies breeding on oranges and those breeding on grapefruit. Within these populations inversions were not distributed at random among individuals; rather there was an observed excess of individuals carrying intermediate numbers, and a deficiency of those carrying very few or very many inversions. While there was no significant linkage disequilibrium associated with this central tendency, there was a significant interchromosomal interaction: flies carrying inversions in chromosome 2 tended not to carry them in chromosome 3, and vice versa.