, the United States and its allies conducted one of the most operationally successful wars in history, a conflict in which air operations played a preeminent role. The Gulf War Air Power Survey was commissioned on 22 August 1 99'. to review all aspects of air warfare in the Persian Gulf for use by the United S~ates Air Force, but it was not to confine itself to discussion of that institution. The Survey has produced reports on planning, the conduct of operations, the effects of the air campaign, command and control, logistics, air base Support,-space, weapons and tactics, as well as a chronology and a compendium of statistics on the war. It has prepared as well a summary report and some shorter papers and assembled an archive composed of paper, microfilm, and electronic records, all of which have beeit deposited at the Air Force Historical Research Agency at Maxwell Air Force Base, Alabama. The Survey was just that, an attempt to provide a comprehensive and documented account of the war. It is not a definitive history: that wifi await the passage of time and the opening of sources (Iraqi records, f-r example) that were niot available to Survey researchers. Nor is it a summary of lessons learned: other organizations, including many within the Air Force, have already done that. Rather, the Survey provides an analytical and evidentiary point of departure for future studies of the air campaign. It concentrates on an analysis of the operational level of war in the belief that this level of warfare is at once one of the most difficult to characterize and one of the most important to understand.
Maternal inheritance of mitochondrial DNA (mtDNA) was originally thought to prevent any response to selection on male phenotypic variation attributable to mtDNA, resulting in a male-biased mtDNA mutation load ("mother's curse"). However, the theory underpinning this claim implicitly assumes that a male's mtDNA has no effect on the fitness of females he comes into contact with. If such "mitochondrially encoded indirect genetics effects" (mtIGEs) do in fact exist, and there is relatedness between the mitochondrial genomes of interacting males and females, male mtDNA-encoded traits can undergo adaptation after all. We tested this possibility using strains of Drosophila melanogaster that differ in their mtDNA. Our experiments indicate that female fitness is influenced by the mtDNA carried by males that the females encounter, which could plausibly allow the mitochondrial genome to evolve via kin selection. We argue that mtIGEs are probably common, and that this might ameliorate or exacerbate mother's curse. K E Y W O R D S indirect genetic effects, interacting phenotypes, kin selection, mtDNA, mutation load, sexual conflict S U PP O RTI N G I N FO R M ATI O N Additional supporting information may be found online in the Supporting Information section. How to cite this article: Keaney TA, Wong HWS, Dowling DK, Jones TM, Holman L. Mother's curse and indirect genetic effects: Do males matter to mitochondrial genome evolution? J
Assuming that fathers never transmit mitochondrial DNA (mtDNA) to their offspring, mitochondrial mutations that affect male fitness are invisible to direct selection on males, leading to an accumulation of male-harming alleles in the mitochondrial genome (mother's curse). However, male phenotypes encoded by mtDNA can still undergo adaptation via kin selection provided that males interact with females carrying related mtDNA, such as their sisters. Here, using experiments with Drosophila melanogaster carrying standardized nuclear DNA but distinct mitochondrial DNA, we test whether the mitochondrial haplotype carried by interacting pairs of larvae affects survival to adulthood, as well as the fitness of the adults. Although mtDNA had no detectable direct or indirect genetic effect on larva-to-adult survival, the fitness of male and female adults was significantly affected by their own mtDNA and the mtDNA carried by their social partner in the larval stage. Thus, mtDNA mutations that alter the effect of male larvae on nearby female larvae (which often carry the same mutation, due to kinship) could theoretically respond to kin selection. We discuss the implications of our findings for the evolution of mitochondria and other maternally inherited endosymbionts.
The novel assay described has potential for application as a rapid, sensitive, high-throughput screening method in the clinical diagnostics setting.
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