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contains no other information than the thermodynamic temperature3. But last year Sanwal et al.4 reported observations using Chandra of the spectrum of the young isolated neutron star 1E1207.45209. The data show two distinct dips in the energy spectrum of photons emitted from the star, centred at 0.7 and 1.4 keV. Here was the first spectral signature of an interaction of the surface radiation with something in the stellar atmosphere but what? These features, known as absorption lines, were difficult to explain. Sanwal et al. speculated that they are due to absorption by singly ionized helium atoms in the stellar atmosphere; others, however, argued that transitions in highly ionized oxygen or neon atoms are the more likely explanation5,6. The energies at which the features appear 0.7 and 1.4 keV are in the ratio 1:2. That suggests another interpretation. Charged particles in a magnetic field oscillate at a particular, resonant frequency this is called cyclotron resonance. For a magnetic field strength of about 61010 gauss, the resonant frequency of electron oscillations corresponds to an absorbed-photon energy of 0.7 keV; absorption will also occur at multiples of this fundamental energy. Protons can resonantly absorb radiation in a magnetic field, but, for a given resonant frequency, the corresponding magnetic field would be 2,000 times higher than in the case of cyclotron resonance of electrons (because a proton is 2,000 times heavier than an electron). From an accurate timing analysis of the spin behaviour of 1E1207.45209, Sanwal et al.4 estimate that the star has a surface magnetic field strength of 231012 gauss. But if the 0.7-keV feature in the X-ray spectrum is a cyclotron resonance, this doesnt add up for an electron resonance, the field strength would be much lower than this value, whereas a proton resonance would require a field strength that is much higher. For this reason, the cyclotron-resonance interpretation was initially rejected. But Bignami et al.2 now report more detailed observations of the same neutron star using cameras onboard XMM-Newton, data that provide convincing proof of the cyclotron interpretation. Bignami et al. have found a third feature at about 2.1 keV and perhaps even a fourth feature around 2.8 keV almost precisely three and four times 0.7 keV, respectively. What makes the identification compelling is that all of the features vary systematically in strength (and perhaps even in shape) with the spin phase of the star, ruling out the possibility of this being a chance conspiracy of noise fluctuations in the data. Cyclotron resonances have been seen before in a handful of neutron stars in binary systems, the first one being Hercules X-1 (ref. 7). But never before has a triple harmonic structure been seen so convincingly in an isolated neutron star.
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The implications of this beautiful spectrum are somewhat puzzling, though. First, there is the obvious question of why only this object shows cyclotron resonances. (There is a recent report8 of a broad feature in the spectrum of the isolated neutron star RBS1223, but even if that also turns out to be a cyclotron resonance, it still means that only a small fraction of isolated neutron stars show these kinds of spectral features.) This question translates into the equivalent statement that only a small fraction of isolated young neutron stars have a low magnetic field strength, one that is in the right range for electron cyclotron resonances to show up in the 0.110-keV band. And there is the problem that the magnetic field strength estimated from the resonance energy is inconsistent with the field strength derived from the stars observed spin rate and deceleration rate. Perhaps there are additional torques on 1E1207.45209. That would clearly have implications for our understanding of the angular momentum and magnetic field strength existing at the birth of a neutron star an understanding that is already under siege on another flank, with the discovery of magnetars. These neutron stars have superstrong magnetic fields9 that exceed a staggering 1014 gauss. Whatever the answer, the fact that 1E1207.45209 shows definite structure in
Evolutionary biology

the spectrum of light emitted near or at its surface is an important finding. Physical clues to the fundamental properties of neutron stars are rare, and this discovery adds an important piece to the puzzle. It will be interesting to see whether the detailed shape of the spectrum can be made to fit a model for the atmosphere, and what the resulting distribution of the field, and perhaps the thermal structure of the atmosphere, looks like. And the discovery itself will undoubtedly add impetus to attempts to find spectral structure in the emissions of other neutron stars. s
Frits Paerels is in the Department of Astronomy and the Columbia Astrophysics Laboratory, Columbia University, New York, New York 10027, USA. e-mail: frits@astro.columbia.edu
1. Miller, C. Nature 420, 3133 (2002). 2. Bignami, G. F., Caraveo, P. A., De Luca, A., & Mereghetti, S. Nature 423, 725727 (2003). 3. Zavlin, V. E. & Pavlov, G. G. in Proc 270. WE-Heraeus Seminar on Neutron Stars, Pulsars, and Supernova Remnants (eds Becker, W., Lesch, H. & Trmper, J.) 263272 (Max Planck Inst. Extraterrestr. Phys., Garching, 2002). 4. Sanwal, D., Pavlov, G. G., Zavlin, V. E. & Teter, M. A. Astrophys. J. 574, L61L64 (2002). 5. Hailey, C. J. & Mori, K. Astrophys. J. 578, L133L136 (2002). 6. Mori, K. & Hailey, C. J. Preprint astro-ph/0301161 at <http://arXiv.org> (2003). 7. Trmper, J. et al. Astrophys. J. 219, L105L110 (1978). 8. Haberl, F., Schwope, A. D., Hambaryan, V., Hasinger, G. & Motch, C. Astron. Astrophys. 403, L19L23 (2003). 9. Duncan, R. C. & Thompson, C. Astrophys. J. 392, L9L13 (1992).

Genes to make new species


Mohamed A. F. Noor A long-term goal of studies of the way in which new species form has been to identify the genes involved, and the forces that drive their evolution. That goal is now being realized and natural selection plays a major part.
or some 70 years, researchers have been crossing different fruitfly species in an attempt to answer one of the most fundamental questions in evolutionary biology: what are the genetic changes that cause one species to split into two? Speciation is known to involve genes that prevent two individuals from mating or, if they do mate, from producing living, fertile offspring. These genes contribute, for instance, to mate discrimination, the success of fertilization, adaptation to particular environments, and the physiological defects of the hybrid offspring that are produced when two species do interbreed. But the identity of such genes, their normal functions and the forces that shaped their evolution are largely unknown. There are two major difficulties in tracking them down. First, species that fail to produce any living or fertile hybrids cannot be studied through genetic crosses. And second, mate discrimination and hybrid sterility often result from the actions of many

genes, sometimes interacting in complex ways. So, even in species for which complete genome sequences are available, identifying the genes that cause speciation has proved difficult. On page 715 of this issue, however, Presgraves and colleagues1 describe how they mapped, identified and analysed one such gene. Found on chromosome 3 of the fruitfly Drosophila simulans, it interacts with one or more unknown genes on the Drosophila melanogaster X chromosome to cause the death of male hybrid offspring. In the two individual species, the protein products of these genes function quite happily together. But the gene on chromosome 3 has evolved divergently in the two species, such that it can no longer function with the X-chromosome genes of the other species in a hybrid male. The authors began their study with fly strains lacking specific small segments of their genomes, and used them to map the hybrid-lethality effect to a single-copy gene
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has been little evidence that genes involved in speciation are necessarily direct targets of natural selection, or are evolving rapidly. Presgraves et al.1 also address this issue. The authors sequenced and analysed 15 copies of Nup96 from different D. simulans and D. melanogaster individuals. Identifying differences in DNA sequences and putative amino-acid sequences both within and between species, they found the signature of positive natural selection early in the lineages of each species. The results show that the death of hybrid males produced by these species evolved at least in part through changes in the sequence of the Nup96 protein, and not through changes in the sequence of genetic regulatory elements. These findings accord with other data suggesting that the characteristics that define species are often the products of natural selection. Many analyses of genes involved in reproduction have identified the signature of natural selection5,6. For instance, sequence differences in the proteins encoded by some of these genes prevent fertilization between sperm and eggs of different species. Similarly, the Drosophila Odysseus gene sequences bear the signature of natural selection3. Finally, the Hybrid male rescue gene, which affects both hybrid viability and female sterility, also shows an extraordinarily large number of differences between D. melanogaster and D. simulans7. It seems from the emerging genetic data, then, that directional natural selection regularly affects genes that contribute to speciation. This is an exciting time for speciation researchers. Studies such as that of Presgraves et al.1 are finally identifying the precise genes that contribute to speciation and the forces that drove their evolution. A decade ago, the joke was that spell-checkers regularly attempted to substitute the word speciation with speculation. Now, the availability of numerous whole-genome sequences, the ability to survey patterns of expression for all genes in a genome simultaneously, and tremendous improvements in computational efficiency are bringing some of the most concrete advances in our understanding of the genetic changes that cause speciation. Speculation in this area will soon be a thing of the past. s
Mohamed A. F. Noor is in the Department of Biological Sciences, Louisiana State University, Baton Rouge, Louisiana 70803, USA. e-mail: mnoor@lsu.edu
1. Presgraves, D. C., Balagopalan, L., Abmayr, S. M. & Orr, H. A. Nature 423, 715719 (2003). 2. Glazier, A. M., Nadeau, J. H. & Aitman, T. J. Science 298, 23452349 (2002). 3. Ting, C.-T., Tsaur, S.-C., Wu, M.-L. & Wu, C.-I. Science 282, 15011504 (1998). 4. Walter, R. B. & Kazianis, S. ILAR J. 42, 299321 (2001). 5. Swanson, W. J. & Vacquier, V. D. Nature Rev. Genet. 3, 137144 (2002). 6. Swanson, W. J. & Vacquier, V. D. Annu. Rev. Ecol. Syst. 33, 161179 (2002). 7. Barbash, D. A., Siino, D. F., Tarone, A. M. & Roote, J. Proc. Natl Acad. Sci. USA 100, 53025307 (2003).
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Figure 1 Subjects of speciation research: Drosophila melanogaster and Xiphophorus maculatus (inset).

on D. simulans chromosome 3. This gene encodes two nucleoporin proteins, Nup96 and Nup98, which are part of a large complex that is needed to shuttle RNAs and proteins between the nucleus and cytoplasm of cells. Presgraves et al. rigorously confirmed their genetic mapping data by complementation testing: they performed individual crosses between D. simulans and strains of D. melanogaster (Fig. 1) that had mutations in each of 12 genes from the chromosomal region that includes the nucleoporin gene. Only those crosses involving D. melanogaster strains that had mutations in the nucleoporin gene failed to produce hybrid males, showing that this gene contributes to the death of hybrid males. Glazier et al.2 have suggested that functional tests, such as complementation tests, are a prerequisite for identifying genes that contribute to complex traits, and Presgraves and colleagues study satisfies this criterion. Presgraves et al. further showed that the D. simulans Nup96 protein and probably one specific end of it, the aminoterminal region bore the sequence variations that caused death when it was associated with the D. melanogaster X chromosome. Two other genes that affect the fitness of hybrids have received much attention recently: the Odysseus gene in Drosophila species, and the Xmrk-2 gene in Xiphophorus fish. Odysseus is thought to contribute to the sterility of hybrid males produced by crossing D. simulans with D. mauritiana3. It was mapped to a chromosomal region bearing a sequence known as a homeobox; genes containing this sequence are usually important in development. But five years after the identification of Odysseus, no functional tests have been
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reported that support its role in hybrid sterility and its functional equivalence to the homeobox gene. So the particular homeobox gene that is purported to be Odysseus is still called OdsH (Odysseus-site homeobox gene), and Presgraves and colleagues demonstration of the effect of the Nup96 gene on hybrid fitness is more compelling. The Xmrk-2 gene, meanwhile, is overexpressed in hybrids produced by Xiphophorus maculatus platyfish and X. helleri swordtails, and causes invasive melanomas that can be fatal4. Functional tests have confirmed the role of this gene in tumour formation but its effects are inconsistent when different strains of platyfish are used. It will be interesting to see whether Nup96 from different strains of D. simulans also has variable effects on hybrid-male viability. The findings could help to determine the particular variations in the Nup96 sequence that cause death. One reason why evolutionary biologists seek specific genes associated with speciation is that, since Darwins time, considerable debate has surrounded the exact role that natural selection plays in species formation. Statistical analyses of species-to-species variations in the sequence of a gene can indicate whether that gene was the target of natural selection: DNA sequences that are subject to recurrent positive natural selection often exhibit more variations that alter the amino-acid sequence they encode than variations that do not. In todays era of genomics, many researchers have identified rapidly evolving genes that appear to be targets of natural selection, and have suggested that these may be important in species formation. Until recently, however, there
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