The Evolutionary History of DROSOPHILA BUZZATII. Xii. the Genetic Basis of Sterility in Hybrids between D. BUZZATII and Its Sibling D. SERIDO from Argentina

The genetic basis of hybrid sterility has been investigated in backcross segmental hybrids between two sibling species, Drosophila buzzatii and D. serido. Asynapsis of homologous bands in hybrid polytene chromosomes has been used to identify the D. serido chromosome segments introgressed into the D. buzzatti genome. All the investigated chromosomes contain male sterility factors. For autosomes, sterility is produced when an introgressed D. serido chromosome segment, or combination of segments, reaches a minimum size. On the other hand, any introgressed X chromosome segment from D. serido, irrespective of its size, produces either male hybrid sterility or inviability.     

Phylogenetic Relationships among DROSOPHILA LONGICORNIS, DROSOPHILA PROPACHUCA and DROSOPHILA PACHUCA, a Triad of Sibling Species

Drosophila longicornis, D. propachuca and D. pachuca comprise a triad of sibling species. They are morphologically indistinguishable, sympatric forms that, under laboratory conditions, are capable of exchanging genes through the production of fertile F₁ females. However, we have no evidence for introgressive hybridization in nature. The chromosomal constitution of our strains indicates that the ancestral species had the Primitive E gene sequence, and therefore differed from the standard repleta sequence by being Xabc; 2abcg; 3abc. This Primitive E sequence is found in both D. propachuca and D. longicornis. Each of these two species has its own unique rearrangements. D. pachuca is a derived species, which evolved from propachuca. It is cytologically more advanced and has, as its most primitive gene arrangement, one of the more advanced arrangements found in propachuca.     

Selective Introgression of Paracentric Inversions between Two Sibling Species of the Anopheles Gambiae Complex

The Anopheles gambiae complex includes the major vectors of malaria in sub-Saharan Africa where >80% of all world-wide cases occur. These mosquitoes are characterized by chromosomal inversions associated to the speciation process and to intraspecific ecological and behavioral flexibility. It has been postulated that introgressive hybridization has selectively transferred inversions on the second chromosome between A. gambiae and A. arabiensis, the two most important vectors of malaria. Here we directly test this hypothesis with laboratory experiments in which hybrid populations were established and the fate of chromosomal inversions were followed. Consistent with the hypothesis, ``foreign' X chromosomes were eliminated within two generations, while some ``foreign' second chromosomes persisted for the duration of the experiments and, judging from the excess of heterozygotes, established stable heterotic polymorphisms. Only those second chromosome inversions found naturally in the species could be introgressed.     

Evolutionary Relationships between Rhynchosporium lolii sp. nov. and Other Rhynchosporium Species on Grasses

The fungal genus Rhynchosporium (causative agent of leaf blotch) contains several host-specialised species, including R. commune (colonising barley and brome-grass), R. agropyri (couch-grass), R. secalis (rye and triticale) and the more distantly related R. orthosporum (cocksfoot). This study used molecular fingerprinting, multilocus DNA sequence data, conidial morphology, host range tests and scanning electron microscopy to investigate the relationship between Rhynchosporium species on ryegrasses, both economically important forage grasses and common wild grasses in many cereal growing areas, and other plant species. Two different types of Rhynchosporium were found on ryegrasses in the UK. Firstly, there were isolates of R. commune that were pathogenic to both barley and Italian ryegrass. Secondly, there were isolates of a new species, here named R. lolii, that were pathogenic only to ryegrass species. R. lolii was most closely related to R. orthosporum, but exhibited clear molecular, morphological and host range differences. The species was estimated to have diverged from R. orthosporum ca. 5735 years before the present. The colonisation strategy of all of the different Rhynchosporium species involved extensive hyphal growth in the sub-cuticular regions of the leaves. Finally, new species-specific PCR diagnostic tests were developed that could distinguish between these five closely related Rhynchosporium species.     

The Evolutionary History of Protein Domains Viewed by Species Phylogeny

Background

Protein structural domains are evolutionary units whose relationships can be detected over long evolutionary distances. The evolutionary history of protein domains, including the origin of protein domains, the identification of domain loss, transfer, duplication and combination with other domains to form new proteins, and the formation of the entire protein domain repertoire, are of great interest.

Methodology/Principal Findings

A methodology is presented for providing a parsimonious domain history based on gain, loss, vertical and horizontal transfer derived from the complete genomic domain assignments of 1015 organisms across the tree of life. When mapped to species trees the evolutionary history of domains and domain combinations is revealed, and the general evolutionary trend of domain and combination is analyzed.

Conclusions/Significance

We show that this approach provides a powerful tool to study how new proteins and functions emerged and to study such processes as horizontal gene transfer among more distant species.     

The Cytogenetic Study of Crossing over in Interspecific Hybrids between DROSOPHILA VIRILIS and D. TEXANA

Spontaneous crossing over was studied by means of combined cytological and genetic methods in F₁ Drosophila virilis x D. texana females (series I) and in D. virilis females carrying a D. texana fifth chromosome in heterozygous condition (series II). The main criterion utilized to distinguish the oogonial crossovers from the meiotic ones is the identity of cytological positions of genetic exchange in crossovers constituting a cluster. Five clusters of crossovers with identical positions of exchange were found in the first series of experiments. In the second series of experiments not a single cluster of crossovers resulting from oogonial crossing over was found.     

Lack of Genic Similarity between Two Sibling Species of Drosophila as Revealed by Varied Techniques

Acrylamide gel electrophoresis was performed on the enzyme xanthine dehydrogenase in sixty isochromosomal lines of Drosophila persimilis from three geographic populations. Sequential electrophoretic analysis using varied gel concentrations and buffers revealed twenty-three alleles in this species where only five had been described previously. These new electrophoretic techniques also detected a profound increase in divergence of gene frequencies at this locus between D. persimilis and its sibling species D. pseudoobscura. The implications of these results for questions of speciation and the maintenance of genetic variability are discussed.     

Genetics of a Difference in Male Cuticular Hydrocarbons between Two Sibling Species, Drosophila Simulans and D. Sechellia

In seven of the eight species of the Drosophila melanogaster group, the predominant cuticular hydrocarbon of males is 7-tricosene, but in the island endemic species D. sechellia it is 6-tricosene. The phylogeny of the group implies that the novel hydrocarbon profile of D. sechellia is a derived character. Genetic analysis of hybrids between D. sechellia and its close relative D. simulans show that each of the five major chromosome arms carries at least one gene affecting the ratio of the two tricosene isomers, with the right arm of the third chromosome having the largest effect. The species difference in this character is therefore polygenic with the effects of the different chromosome arms generally additive, although there is some epistasis among third-chromosome genes. Observations of courtship by males who have been coated with foreign hydrocarbons suggest that a male's hydrocarbon profile may slightly affect the degree of sexual isolation in one of the reciprocal hybridizations between these species, but that this role is small compared to that played by hydrocarbon differences between females.     

The Broom of the Sorcerer''s Apprentice: The Fine Structure of a Chromosomal Region Causing Reproductive Isolation between Two Sibling Species of Drosophila

How many genes contribute to reproductive isolation between closely related species? We determined the number of genes located in the 9D-12B region of the Drosophila mauritiana X chromosome that cause hybrid male sterility in a D. simulans background. Previous low resolution studies suggested that a single hybrid sterility factor was associated with this region. In this study, by taking advantage of a cluster of visible and DNA markers, we identified three D. mauritiana factors in this region and then subjected one of them to detailed analysis. This factor again turned out to be comprised of three factors; one of which, mapped to within 200 kb, may in fact be two factors. The title refers to this exercise of splitting sterile introgressions into ever smaller ones, each of which retains partial or full sterility effects. In a region representing a mere 3% of the Drosophila genome, no fewer than six loci of hybrid sterility were identified between two sibling species that have not shown clear divergence at the molecular level. These results suggest that levels of genetic divergence between closely related species may be quite high for functionally important traits even when the opposite is true for randomly chosen loci.     

Evaluation of Phylogenetic Relationships between Five Polyploid Aegilops L. Species of the U-Genome Cluster by Means of Chromosome Analysis

Four tetraploid (Aegilops ovata, Ae. biuncialis, Ae. columnaris, and Ae. triaristata) and one hexaploid (Ae. recta) species of the U-genome cluster were studied using C-banding technique. All species displayed broad C-banding polymorphism and high frequency of chromosomal rearrangements. Chromosomal rearrangements were represented by paracentric inversions and intragenomic and intergenomic translocations. We found that the processes of intraspecific divergence of Ae. ovata, Ae. biuncialis,and Ae. columnaris were probably associated with introgression of genetic material from other species. The results obtained confirmed that tetraploid species Ae. ovata and Ae. biuncialis occurred as a result of hybridization of a diploidAe. umbellulata with Ae. comosa and Ae. heldreichii, respectively. The dissimilarity of the C-banding patterns of several chromosomes of these tetraploid species and their ancestral diploid forms indicated that chromosomal aberrations might have taken place during their speciation. Significant differences of karyotype structure, total amount and distribution of C-heterochromatin found between Ae. columnarisand Ae. triaristata, on the one hand, and Ae. ovata and Ae. biuncialis, on the other, evidenced in favor of different origin of these groups of species. In turn, similarity of the C-banding patterns of Ae. columnaris and Ae. triaristata chromosomes suggested that they were derived from a common ancestor. A diploid species Ae. umbellulata was the U-genome donor of Ae. columnaris and Ae. triaristata;however, the donor of the second genome of these species was not determined. We assumed that these tetraploid species occurred as a result of introgressive hybridization. Similarity of the C-banding patterns of chromosomes of Ae. recta and its parental species Ae. triaristata and Ae. uniaristata indicated that the formation of the hexaploid form was not associated with large modifications of the parental genomes.     

The Chromosomal Basis of Sexual Isolation in Two Sibling Species of Drosophila: D. ARIZONENSIS and D. MOJAVENSIS

The chromosomal determination of interspecific differences in mating behavior was studied in the interfertile pair, Drosophila arizonensis and Drosophila mojavensis, by means of chromosomal substitutions. Interspecific crossing over was avoided by crossing hybrid males to parental females, and identification of the origin of each chromosome in backcrossed hybrids was possible by means of allozyme markers. It was found that male mating behavior is controlled by factors located in the PGM-marked chromosome (which, in other Drosophila species, is part of the X chromosome) and in the Y chromosome. The other chromosomes influence male sexual behavior through their interactions with each other and with the PGM-marked chromosome, but their overall effect is minor. Female mating behavior is controlled by factors located in the ODH-marked and AMY-marked chromosomes, with the other chromosomes exercising a small additive effect. Hence, the two sex-specific behaviors are under different genetic control. Cytoplasmic origin has no effect on the mating behavior of either sex. There appears to be no correlation between a chromosome's structural diversity (i.e., amounts of inversion polymorphism within a species or numbers of fixed inversions across species) and its contribution to sexual isolation. These findings are in general agreement with those from similar Drosophila studies and may not be specific to the species studied here.     

The Evolutionary History of the Transposable Element Penelope in the Drosophila virilis Group of Species

We have used phylogenetic techniques to study the evolutionary history of the Penelope transposable element in the Drosophila virilis species group. Two divergent types of Penelope have been detected, one previously described, clade I, and a new one which we have termed clade III. The phylogeny of some copies of the Penelope clade I element was partially consistent with the species phylogeny of the D. montana subphylad, suggesting cospeciation and allowing the estimation of the evolutionary rate of Penelope. Divergence times of elements found in different species are younger than the age of the species, suggesting horizontal transfer events. Electronic Supplementary Material Electronic Supplementary material is available for this article at and accessible for authorised users. [Reviewing Editor: Dr. Dmitri Petrov]     

Strong Purifying Selection on the Odysseus Gene in Two Clades of Sibling Species of the Drosophila montium Species Subgroup

The Odysseus (OdsH) gene was duplicated from its ancestral neuron-expressed gene, unc-4, and then evolved very rapidly under strong positive Darwinian selection as a speciation gene causing hybrid-male sterility between closely related species of the Drosophila simulans clade. Has OdsH also experienced similar positive selection between Drosophila sibling species other than those of the simulans clade? We cloned and sequenced OdsH and unc-4 from two clades of the Drosophila montium species subgroup, the Drosophila lini and the Drosophila kikkawai clades. The ratios of Ka/Ks for OdsH were remarkably low between sibling species of these two clades, suggesting that OdsH has been subjected to strong purifying selection in these two clades.     

The Evolutionary History of the Australopiths

The australopiths are a group of early hominins (humans and their close extinct relatives) that lived in Africa between approximately 4.1 and 1.4 million years ago. Formerly known as the australopithecines, they are not a “natural” group, in that they do not represent all of the descendants of a single common ancestor (i.e., they are not a “clade”). Rather, they are grouped together informally because nearly all share a similar adaptive grade (i.e., they have similar adaptations). In particular, they are bipedal apes that, to a greater or lesser extent, exhibit enlarged molar and premolar teeth (postcanine megadontia) and other associated modifications to their feeding apparatuses. Dietary adaptations clearly played an important role in shaping their evolutionary history. They also are distinguished by their lack of derived features typically associated with the genus Homo, such as a large brain, a broad complement of adaptations for manual dexterity, and advanced tool use. However, Homo is almost certainly descended from an australopith ancestor, so at least one or some australopiths belong directly to the human lineage. Regardless, australopiths had a rich evolutionary history deserving of study independent of questions about our direct ancestry. They were diverse, geographically widespread, and anatomically derived, they lived through periods of pronounced climate change, and their story dominates the narrative of human evolution for millions of years.     

The Phylogenetic Relationships of the Members of the DROSOPHILA ROBUSTA Group

The phylogenetic relationships among the species of the D. robusta group were investigated by the analysis of chromosomal differences. Six of the ten known members of the D. robusta group were available for the study: D. colorata and D. robusta from the United States, and D. sordidula, D. pseudosordidula, D. lacertosa, and D. moriwakii from Japan. Analysis of the metaphase chromosomes from larval ganglion cells suggests that D. moriwakii and D. colorata, with rod-shaped X-chromosomes, are the more ancestral species, while D. sordidula, D. pseudosordidula, D. robusta, and D. lacertosa, with V-shaped X-chromosomes, are derived. The ancestral position of D. colorata and D. moriwakii is further strengthened by the fact that these are the two species in the D. robusta group that are cytologically closest to D. nigromelanica of the related D. melanica group. Of the four derived species, D. sordidula was found to be the closest to the ancestral species. The phylogeny based on the analysis of the gene sequences in the homologous chromosomes agreed with that indicated by the metaphase chromosomes. Since all attempts to obtain hybrids were unsuccessful except for the cross involving D. moriwakii females and D. colorata males, photographic maps of the salivary chromosomes were used to determine homology between the chromosomes of the different species. Evidence is presented to indicate that the D. robusta group originated in Asia (Japan), and that there were two migrations to the New World, the first leading to D. robusta, and the second to D. colorata. It is suggested that the route of migrations was across the Bering Land Bridge, and further, that the migrations occurred during the period from late Oligocene to middle Miocene, 20-25 million years ago.     

A Genetic Basis for the Inviability of Hybrids between Sibling Species of Drosophila

A mutation of Drosophila melanogaster whose only known effect is the rescue of otherwise lethal interspecific hybrids has been characterized. This mutation, Hmr, maps to 1-31.84 (9D1-9E4). Hmr may be the consequence of a P element insertion. It rescues hybrid males from the cross of D. melanogaster females to males of its three sibling species, D. simulans, D. mauritiana and D. sechellia. This rescue is recessive, since hybrid males that carry both Hmr and a duplication expected to be Hmr+ are not rescued. Hmr also rescues the otherwise inviable female hybrids from the cross of compound-X D. melanogaster females to males of its sibling species. This rescue is also recessive, since a compound-X heterozygous for Hmr does not rescue. Another mutation, discovered on the In(1)AB chromosome of D. melanogaster, is also found to rescue normally inviable species hybrids: unlike Hmr, however, In(1)AB rescues hybrid females from the cross of In(1)AB/Y males to sibling females, as well as hybrid males from the cross of In(1)AB females to sibling males. These data are interpreted on the basis of a model for the genetic basis of hybrid inviability of complementary genes.     

Typology Reconsidered: Two Doctrines on the History of Evolutionary Biology

Recent historiography of 19th century biology supports the revision of two traditional doctrines about the history of biology. First, the most important and widespread biological debate around the time of Darwin was not evolution versus creation, but biological functionalism versus structuralism. Second, the idealist and typological structuralist theories of the time were not particularly anti-evolutionary. Typological theories provided argumentation and evidence that was crucial to the refutation of Natural Theological creationism. The contrast between functionalist and structuralist approaches to biology continues today, and the historical misunderstanding of 19th century typological biology may be one of its effects. This historical case can shed light on current controversies regarding the relevance of developmental biology to evolution.