The multisubunit MSL dosage compensation complex binds to hundreds of sites along the Drosophila single male X chromosome, mediating its hypertranscription. The male X chromosome is also coated with noncoding roX RNAs. When either msl3, mle, or mof is mutant, a partial MSL complex is bound at only approximately 35 unusual sites distributed along the X. We show that two of these sites are the roX1 and roX2 genes and postulate that one of their functions is to provide entry sites for the MSL complex to recognize the X chromosome. The roX1 gene provides a nucleation site for extensive spreading of the MSL complex into flanking chromatin even when moved to an autosome. The spreading can occur in cis or in trans between paired homologs. We present a model for how the dosage compensation complex recognizes X chromatin.
A high spectral resolution observation of the diffuse X-ray background in the 60 -1000 eV energy range has been made using an array of thirty-six 1 mm 2 microcalorimeters flown on a sounding rocket. Detector energy resolution ranged from 5-12 eV FWHM, and a composite spectrum of 1 steradian of the background centered at l = 90°, b = +60° was obtained with a net resolution of ~ 9 eV. The target area includes bright 1/4 keV regions, but avoids Loop I and the North Polar Spur. Lines of C VI, O VII, and O VIII are clearly detected with intensities of 5.4 ± 2.3, 4.8 ± 0.8, and 1.6 ± 0.4 photons cm -2 s -1 sr -1 , respectively. The oxygen lines alone account for a majority of the diffuse background observed in the ROSAT R4 band that is not due to resolved extragalactic discrete sources. We also have a positive detection of the Fe-M line complex near 70 eV at an intensity consistent with previous upper limits that indicate substantial gas phase depletion of iron. We include a detailed description of the instrument and its detectors.
The energy splitting of the 229Th ground-state doublet is measured to be 7.6+/-0.5 eV, significantly greater than earlier measurements. Gamma rays produced following the alpha decay of 233U (105 muCi) were counted in the NASA/electron beam ion trap x-ray microcalorimeter spectrometer with an experimental energy resolution of 26 eV (FWHM). A difference technique was applied to the gamma-ray decay of the 71.82 keV level that populates both members of the doublet. A positive correction amounting to 0.6 eV was made for the unobserved interband decay of the 29.19 keV state (29.19-->0 keV).
Male-specific lethal-2 (msl-2) is a RING finger protein that is required for X chromosome dosage compensation in Drosophila males. Consistent with the formation of a dosage compensation protein complex, msl-2 colocalizes with the other MSL proteins on the male X chromosome and coimmunoprecipitates with msl-1 from male larval extracts. Ectopic expression of msl-2 in females results in the appearance of the other MSL dosage compensation regulators on the female X chromosomes and decreased female viability. We suggest that msl-2 RNA is the primary target of SxI regulation in the dosage compensation pathway and present a speculative model for the regulation of two distinct modes of dosage compensation by SxI.
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