The Life Cycle P Drosophila Lifecycle.

Neuroptera Coleoptera Hymenoptera Lepidoptera Mecoptera Siphonoptera Strepsiptera Diptera ~250 Mya

Nematocera Brachycera Muscomorpha Cyclorrhapha Schizophora Calyptrata Acalyptrata

Camillidae Steganinae Cladocheata etc Zaprionus etc s.g. Drosophila Chymomyza etc s.g. Sophopora

Drosophila melanogaster subgroup Drosophila melanogaster - cosmopolitan - 1830 Drosophila simulans - cosmopolitan - 1919 Drosophila mauritiana - Mauritius - 1974 Drosophila sechellia - Seychelles -1981 Drosophila yakuba - Equatorial Africa - 1954 Drosophila santomea - Sao Tome - 2000 Drosophila teissieri - Equatorial Africa - 1979 Drosophila erecta - Central West Africa - 1974 Drosophila orena - Cameroons - 1978

mel f x sim m sim f x mel m Viable, sterile f Viable, sterile m Larval lethal m Embryo lethal f aabb AAbb x aaBB AaBb Hmr allele of melanogaster (transcription factor) Lhr allele of simulans (chromatin binding protein)

“Eventually, the story of the chromosomal mechanisms and its evolution will have to be entirely rewritten in molecular terms.” Michael White, 1973, Animal Cytology & Evolution, 2nd edition.

Paracentric inversions are the most common rearrangement Segregating in 106 out 183 species. - 57% - (Powell 1997). 22000-56000 inversions have become fixed during the evolution of the genus (Clayton & Guest 1986).

a b c d e f g h i j k l m n o p q r s t a b j i h g f e d c k l m n o p q r s t a b j l o n m l k c d e f h i p q r s t a b c d e f g h i o n m l k j p q r s t a b k l m n o i h g f e d c j p q r s t

12 genomes have been sequenced in the genus Drosophila

1. 2. 3. Ectopic recombination between homologous fragments of DNA ( Cáceres et al. Science 1999 )

3R of D. melanogaster 3R of D. simulans ~450 Kb ~280 Kb 84F1 93F6-7 BACs BACR16N15 BACR42I20 BACR08K01 BACR07M14 BACR45A07 3R of D. melanogaster centromere telomere 84F1 93F6-7 centromere telomere 3R of D. simulans

Inversion-mediated duplications can result from staggered isochromatid breaks & repair. F G H I J K 5' 3' A B C D E 5' 3' 3' 5' 3' 5' 5' 3' 3' 5' 5' 3' 5' 3' 3' 5' A B C D I H G F E D C B I J K 5' 3' 5' 3' 3' 5' 3' 5' 3' 5'

Lemeunier & Ashburner, 1976

BLAST-N of D. mel 3R transcripts against D. yak 3R J. Ranz, C. Bergman & M. Ashburner

Flanking duplications are a common by-product of the genome reorganization between D. melanogaster and D. yakuba. 20 9 with without

Most inversions occurred in the D. yakuba lineage 1 ( 3.4 % ) 28 ( 96.6 % ) D. melanogaster D. yakuba outgroup species 1: A-B A]-----[B A-B 2: A]-----[B A-B A-B

Large-scale comparison of the gene order between D. melanogaster and D. yakuba. 55 breakpoints = 12 + 22 + 9 + 12 29 inversions Breakpoints Mb Myr

Cluster gram based on the number of breakpoints that correspond to inversions

Genes within genes within genes

Number of antisense/sense overlapping pairs of genes in D. melanogaster

D. melanogaster is different from H. sapiens. RIKEN Group and FANTOM Consortium, Antisense transcription in the mammalian transcriptome. Science 2005 309(5740)

Casey Bergman (Manchester). Marcin von Grotthuss (Cambridge). Thanks to: Jose Ranz (Cambridge). Casey Bergman (Manchester). Marcin von Grotthuss (Cambridge). Karen Eilbeck & Suzi Lewis (Berkeley). Lincoln Stein (CSGL) & Richard Durbin (WTGC) Hadi Quesneville (Paris). The MRC for 25 years of continuous funding. The BBSRC for a grant to Jose. EMBO for support to Marcin. The Royal Society for a Fellowship to Casey.

Annotation Depth of knowledge Breadth of knowledge Detailed analysis (typically biological) of single genes Annotated genome Depth of knowledge Large-scale analysis (typically computational) of entire genome Breadth of knowledge

Assemblies from the 12 ongoing Drosophila sequencing projects are available right now