Evolution of the LINE-like I element in the Drosophila melanogaster species subgroup

LINE-like retrotransposons, the so-called I elements, control the system of I-R (inducer-reactive) hybrid dysgenesis in Drosophila melanogaster. I elements are present in many Drosophila species. It has been suggested that active, complete I elements, located at different sites on the chromosomes, invaded natural populations of D. melanogaster recently (1920–1970). But old strains lacking active I elements have only defective I elements located in the chromocenter. We have cloned I elements from D. melanogaster and the melanogaster subgroup. In D. melanogaster, the nucleotide sequences of chromocentral I elements differed from those on chromosome arms by as much as 7%. All the I elements of D. mauritiana and D. sechellia are more closely related to the chromosomal I elements of D. melanogaster than to the chromocentral I elements in any species. No sequence difference was observed in the surveyed region between two chromosomal I elements isolated from D. melanogaster and one from D. simulans. These findings strongly support the idea that the defective chromocentral I elements of D. melanogaster originated before the species diverged and the chromosomal I elements were eliminated. The chromosomal I elements reinvaded natural populations of D. melanogaster recently, and were possibly introduced from D. simulans by horizontal transmission.     

Evolutionary relationships to parasitism by seven species of the Drosophila melanogaster subgroup

Parasitic wasps are an important component of the niche of Drosophila species. The susceptibility to the Cynipid Leptopilina boulardi was estimated in the seven sibling species of Drosophila belonging to the melanogaster subgroup. Three categories of Hies can be distinguished, according to the level of cellular immune reaction and success of parasitism. Drosophila melanogaster and D. mauritiana belong to the category 1, specified by no encapsulative reaction and a high rate of successful parasitism. Category 2, characterized by a moderate encapsulation rate and a high mortality include D. simulans.5, D. erecta and D. orena. Category 3, with D. yakuba and D. tcissien, is specified by a very low rate or absence of successful parasitism due to a highly efficient immune cellular reaction. This classification parallels the phylogenic relationship based upon polytene chromosome banding sequences. Such specific ditferences in susceptibility to parasites may plan an important role in the competition between these species in Africa.     

Evolutionary relationships to parasitism by seven species of the Drosophila melanogaster subgroup

Parasitic wasps are an important component of the niche of Drosophila species. The susceptibility to the Cynipid Leptopilina boulardi was estimated in the seven sibling species of Drosophila belonging to the melanogaster subgroup. Three categories of Hies can be distinguished, according to the level of cellular immune reaction and success of parasitism. Drosophila melanogaster and D. mauritiana belong to the category 1, specified by no encapsulative reaction and a high rate of successful parasitism. Category 2, characterized by a moderate encapsulation rate and a high mortality include D. simulans.5, D. erecta and D. orena . Category 3, with D. yakuba and D. tcissien , is specified by a very low rate or absence of successful parasitism due to a highly efficient immune cellular reaction. This classification parallels the phylogenic relationship based upon polytene chromosome banding sequences. Such specific ditferences in susceptibility to parasites may plan an important role in the competition between these species in Africa.     

Evolution of eye morphology and rhodopsin expression in the Drosophila melanogaster species subgroup

A striking diversity of compound eye size and shape has evolved among insects. The number of ommatidia and their size are major determinants of the visual sensitivity and acuity of the compound eye. Each ommatidium is composed of eight photoreceptor cells that facilitate the discrimination of different colours via the expression of various light sensitive Rhodopsin proteins. It follows that variation in eye size, shape, and opsin composition is likely to directly influence vision. We analyzed variation in these three traits in D. melanogaster, D. simulans and D. mauritiana. We show that D. mauritiana generally has larger eyes than its sibling species, which is due to a combination of larger ommatidia and more ommatidia. In addition, intra- and inter-specific differences in eye size among D. simulans and D. melanogaster strains are mainly caused by variation in ommatidia number. By applying a geometric morphometrics approach to assess whether the formation of larger eyes influences other parts of the head capsule, we found that an increase in eye size is associated with a reduction in the adjacent face cuticle. Our shape analysis also demonstrates that D. mauritiana eyes are specifically enlarged in the dorsal region. Intriguingly, this dorsal enlargement is associated with enhanced expression of rhodopsin 3 in D. mauritiana. In summary, our data suggests that the morphology and functional properties of the compound eyes vary considerably within and among these closely related Drosophila species and may be part of coordinated morphological changes affecting the head capsule.     

Relationships within the melanogaster species subgroup of the genus Drosophila (Sophophora)

The polytene chromosomes of two new species of Drosophila, D. sechellia and D. orena, both members of the melanogaster species subgroup, are described. The chromosomes of D. sechellia, a species endemic to certain islands in the Seychelles, are homosequential with those of D. simulans and D. mauritiana. The chromosomes of D. orena, a species from the mountains of west Africa, are very similar to those of D. erecta. We discuss the interrelationships of the eight known species of the melanogaster species subgroup, based upon an analysis of their chromosome banding patterns.     

Molecular evolution of the histone 3 multigene family in the Drosophila melanogaster species subgroup

Molecular evolution of the histone multigene family was studied by cloning and sequencing regions of the histone 3 gene in the Drosophila melanogaster species subgroup. Analysis of the nucleotide substitution pattern showed that in the coding region synonymous changes occurred more frequently to A or T in contrast to the GC-rich base composition, while in the 3' region the nucleotide substitutions were most likely in equilibrium. These results suggested that the base composition at the third codon position of the H3 gene, i.e., codon usage, has been changing to A or T in the Drosophila melanogaster species subgroup.     

Divergence and heterogeneity of the histone gene repeating units in the Drosophila melanogaster species subgroup

The repeating units of the histone gene cluster containing the H1, H2A, H2B and H4 genes were amplified by PCR from the Drosophila melanogaster species subgroup, i.e., D. yakuba, D. erecta, D. sechellia, D. mauritiana, D. teissieri and D. orena. The PCR products were cloned and their nucleotide sequences of about 4.6-4.8kbp were determined to elucidate the mechanism of molecular evolution of the histone gene family. The heterogeneity among the histone gene repeating units was 0.6% and 0.7% for D. yakuba and D. sechellia, respectively, indicating the same level of heterogeneity as in the H3 gene region of D. melanogaster. Divergence of the genes among species even in the most closely related ones was much greater than the heterogeneity among family members, indicating a concerted mode of evolution for the histone gene repeating units. Among the species in the D. melanogaster species subgroup, the histone gene regions as well as 3rd codon position of the coding region showed nearly the same GC contents. These results suggested that the previous conclusion on analysis of the H3 gene regions, the gene family evolution in a concerted fashion, holds true for the whole histone gene repeating unit.     

Light dependency of the mating process in Closterium acerosum

Sexual cell division and activation of gametangial cells forconjugation in Closterium acerosum were induced by light. L₂₀₀cells conjugated at maximum level under the following conditions;(i) a light intensity higher than 1,000 lux in a 16-hr lightand 8-hr dark regime and (ii) an illumination time longer than12 hr at 3,000 lux. L₂₀₀ cells also conjugated under continuousillumination at 3,000 lux. The action spectrum for the activation of gametangial cellshad peaks around 450, 611 and 665 nm. 3-(4'-Chlorophenyl)-l,l-dimethylurea (CMU) inhibited the accumulationof carbohydrates and sexual cell division at 10^–5 M andthe activation of gametangial cells for conjugation at 10^–4M. (Received August 15, 1977; )     

The distribution of satellite and main-band DNA components in the melanogaster species subgroup of Drosophila

Fractionation of total adult DNA of five of the seven species of the melanogaster species sub-group of Drosophila in actinomycin D and distamycin A caesium density gradients has revealed the presence of three main-band DNA components, common to all species, and ten satellite DNAs that are distributed between the species. Satellite DNAs are either unique to a species or common to two or more species. The abundance of a common satellite DNA varies between species. There is no simple relationship between the presence of a satellite DNA and a branch point of phylogenetic divergence; nevertheless the arrangement of the species in a phylogeny that is based on the numbers of satellites held in common accurately reflects the pattern of relationships between the same species based on differences in inversions of polytene chromosomes. The species can be similarly arranged according to the compositions of their mitochondrial DNAs. It is possible that the same basic set of sequences, each of low frequency, is common to all species with arbitrary or selected amplification of particular sequences to differing extents in individual species. The conservation of satellites in the group and the close parallel between the distributions of satellites and inversions between the species suggests that either the processes that operate to change both chromosomal phenomena are similarly time-dependent and occurring at relatively low rates or that their rates of change are restricted according to some undetermined functions of these aspects of the genome.     

Evolution of the olfactory code in the Drosophila melanogaster subgroup

The Drosophila melanogaster subgroup has been the focus of numerous studies about evolution. We address the question of how the olfactory code has evolved among the nine sister species. By using in vivo electrophysiological measurements, so called single-cell recordings, we have established the ligand affinity of a defined subset of olfactory receptor neurons (ORNs) across all nine species. We show that the olfactory code as relayed by the investigated subset of ORNs is conserved to a striking degree. Distinct shifts in the code have occurred only within the simulans clade. However, these shifts are restricted to an altered tuning profile of the same single ORN type in all three of the simulans siblings and a more drastic change unique to D. sechellia, involving a complete loss of one sensillum type in favour of another. The alterations observed in D. sechellia may represent a novel host-specific adaptation to its sole host, morinda fruit (Morinda citrifolia). The overall high degree of similarity of the code within the subgroup is intriguing when considering the great variety in distributions as well as in habitat and host choice of the siblings, factors that could greatly affect the olfactory system.     

An intron loss of Dfak gene in species of the Drosophila melanogaster subgroup and phylogenetic analysis

Drosophila focal adhesion kinase (Dfak) gene is a single-copy nuclear gene. Previous study revealed that Drosophila melanogaster and Drosophila simulans had lost an intron precisely within the tyrosine kinase (TyK) domain of this gene. However, this did not happen in several other Drosophila species, including Drosophila elegans, Drosophila ficusphila, Drosophila biarmipes, Drosophila jambulina, Drosophila prostipennis, Drosophila takahashii, and Drosophila pseudoobscura. In the current study, homologous sequences of Drosophila sechellia, Drosophila mauritiana, Drosophila yakuba, Drosophila teissieri, Drosophila santomea, and Drosophila erecta were amplified by polymerase chain reaction, and further sequencing analysis indicated that these species were missing a TyK domain intron, indicating they were closely related. The relationship of the D. melanogaster species group was reconstructed using TyK domain nucleotide sequences. The resulting phylogenetic tree revealed that these 8 species were the most related species in the melanogaster group. These results strongly support previously proposed classifications based on morphological and molecular data.