EPR of Cu\u3csup\u3e2+\u3c/sup\u3e Prion Protein Constructs at 2 GHz Using the \u3cem\u3eg\u3c/em\u3e\u3csub\u3e⊥\u3c/sub\u3e Region to Characterize Nitrogen Ligation

A double octarepeat prion protein construct, which has two histidines, mixed with copper sulfate in a 3:2 molar ratio provides at most three imidazole ligands to each copper ion to form a square-planar Cu2+ complex. This work is concerned with identification of the fourth ligand. A new (to our knowledge) electron paramagnetic resonance method based on analysis of the intense features of the electron paramagnetic resonance spectrum in the g⊥ region at 2 GHz is introduced to distinguish between three and four nitrogen ligands. The methodology was established by studies of a model system consisting of histidine imidazole ligation to Cu2+. In this spectral region at 2 GHz (S-band), g-strain and broadening from the possible rhombic character of the Zeeman interaction are small. The most intense line is identified with the MI = +1/2 extra absorption peak. Spectral simulation demonstrated that this peak is insensitive to cupric Ax and Ay hyperfine interaction. The spectral region to the high-field side of this peak is uncluttered and suitable for analysis of nitrogen superhyperfine couplings to determine the number of nitrogens. The spectral region to the low-field side of the intense extra absorption peak in the g⊥ part of the spectrum is sensitive to the rhombic distortion parameters Ax and Ay. Application of the method to the prion protein system indicates that two species are present and that the dominant species contains four nitrogen ligands. A new loop-gap microwave resonator is described that contains ∼1 mL of frozen sample

Temporal Variability and cooperative Breeding: Testing the Bet-Hedging Hypothesis in the Acorn Woodpecker

Cooperative breeding is generally considered an adaptation to ecological constraints on dispersal and independent breeding, usually due to limited breeding opportunities. Although benefits of cooperative breeding are typically thought of in terms of increased mean reproductive success, it has recently been proposed that this phenomenonmay be a bet-hedging strategy that reduces variance in reproductive success (fecundity variance) inpopulations living inhighly variable environments. We tested this hypothesis using long-term data on the polygynandrous acorn woodpecker (Melanerpes formicivorus). In general, fecundity variance decreased with increasing sociality, at least when controlling for annual variation in ecological conditions. Nonetheless, decreased fecundity variance was insufficient to compensate for reduced per capita reproductive success of larger,more social groups, which typically suffered lower estimatedmean fitness.We did, however, find evidence that sociality in the form of larger group size resulted in increased fitness in years following a small acorn crop due to reduced fecundity variance. Bet-hedging, although not the factor driving sociality in general, may play a role in driving acorn woodpecker group living when acorns are scarce and ecological conditions are poor

The Criticality of Defense-Focused Technical Education

The United States Air Force is a service born of technology, and throughout its history, technology has remained central to its identity and power. From the start, visionary leaders realized the importance of technologically focused education to advancing airpower. Consequently, through the years, institutions of higher learning such as the Air Force Institute of Technology AFIT, as well as the civilian institution program it administers, have continued the meaningful work of developing the technology and organic human capital to sustain the Air Forces edge as a fighting force. As advances in technology have led the Air Force into the new domains and challenges of space and cyberspace, the role of delivering defenseoriented technical education has become even more critical. In this process, leveraging our network of science and technology partners to produce technically educated and operationally focused Airmen has proved as significant as the advances themselves. Because demand for these graduates continues to increase, deliberate investment in science, technology, engineering, and mathematics STEM education must also increase. Today, as yesterday, experienced Air Force leaders with a defense-focused technical education are essential to maintaining our military supremacy, and AFIT continues to meet that need--as it has since its inception in 1919

Variable Helper Effects, Ecological Conditions, and the Evolution of Cooperative Breeding in the Acorn Woodpecker

The ecological conditions leading to delayed dispersal and helping behavior are generally thought to follow one of two contrasting scenarios: that conditions are stable and predictable, resulting in young being ecologically forced to remain as helpers (extrinsic constraints and the habitat saturation hypothesis), or that conditions are highly variable and unpredictable, leading to the need for helpers to raise young, at least when conditions are poor (intrinsic constraints and the hard life hypothesis). We investigated how variability in ecological conditions influences the degree to which helpers augment breeder fitness in the cooperatively breeding acorn woodpecker (Melanerpes formicivorus), a species in which the acorn crop, territory quality, and prior breeding experience all vary in ways that have important effects on fitness. We found that the relationship between ecological conditions and the probability that birds would remain as helpers was variable but that helpers generally yielded greater fitness benefits when ecological conditions were favorable, rather than unfavorable, for breeding. These results affirm the importance of extrinsic constraints to delayed dispersal and cooperative breeding in this species, despite its dependence on a highly variable and unpredictable acorn crop. Our findings also confirm that helpers can have very different fitness effects, depending on conditions, but that those effects are not necessarily greater when breeding conditions are unfavorable

Cooperative Breeding and Long-Distance Dispersal: A Test Using Vagrant Records

Cooperative breeding is generally associated with increased philopatry and sedentariness, presumably because short-distance dispersal facilitates the maintenance of kin groups. There are, however, few data on long-distance dispersal in cooperative breeders-the variable likely to be important for genetic diversification and speciation. We tested the hypothesis that cooperative breeders are less likely to engage in long-distance dispersal events by comparing records of vagrants outside their normal geographic range for matched pairs (cooperatively vs. non-cooperatively breeding) of North American species of birds. Results failed to support the hypothesis of reduced long-distance dispersal among cooperative breeders. Thus, our results counter the conclusion that the lower rate of speciation among cooperative breeding taxa found in recent analyses is a consequence of reduced vagility

Testing Alternative Hypotheses for the Cause of Population Declines: The Case of the Red-Headed Woodpecker

The Red-headed Woodpecker (Melanerpes erythrocephalus) has experienced strong population declines during the past 3 decades. Using North American Breeding Bird Survey (BBS) and Audubon Christmas Bird Count (CBC) data, we investigated 4 hypotheses that may explain this decline, including: (1) interspecific competition with native Red-bellied Woodpeckers (Melanerpes carolinus) and nonnative European Starlings (Sturnus vulgaris); (2) predation by Cooper\u27s Hawks (Accipiter cooperii) and Sharp-shinned Hawks (Accipiter striatus); (3) climate change; and (4) changes in forested area within their range. In analyses of both the breeding and overwintering periods, our results indicated a role of increased accipiter populations in driving Red-headed Woodpecker declines through increased predation. We also found evidence for significant effects of warmer winter temperatures and increased forest cover, both directly and indirectly through their effects on enhancing accipiter populations. In contrast, our results failed to support the hypothesis that interspecific competition with either Red-bellied Woodpeckers or European Starlings has played a role in Red-headed Woodpecker declines. Despite considerable evidence for nest-site competition and aggression between Red-headed Woodpeckers and both Red-bellied Woodpeckers and European Starlings, these interactions do not appear to be limiting Red-headed Woodpecker populations

Habitat Saturation Results in Joint-Nesting Female Coalitions in a Social Bird

Joint nesting by females and cooperative polyandry—cooperatively breeding groups with a male-biased breeder sex ratio—are little-understood, rare breeding systems. We tested alternative hypotheses of factors potentially driving these phenomena in a population of joint-nesting acorn woodpeckers (Melanerpes formicivorus). During periods of high population density and thus low independent breeding opportunities, acorn woodpecker females formed joint-nesting coalitions with close kin. Coalitions were typically associated with groups with a male bias. We found strong evidence for both inter- and intrasexual conflict, as joint nesting conferred a fitness benefit to some males, a significant fitness cost to females, and no gain in per capita reproductive output for either sex. Such conflict, particularly the cost to females, may be an important reason why joint nesting is rare among cooperatively breeding taxa

Territory Inheritance and the Evolution of Cooperative Breeding in the Acorn Woodpecker

There are two main hypotheses for why offspring in cooperatively breeding taxa delay dispersal and remain on their natal territory rather than disperse. First, ecological constraints may force offspring to remain on their natal territory until a reproductive opportunity presents itself in an otherwise saturated habitat. Alternatively, delaying dispersal and helping kin may increase an offspring\u27s inclusive fitness. One means by which offspring might enhance their direct fitness by delaying dispersal is by inheriting breeding status on their natal territory. Such territory inheritance regularly occurs in acorn woodpeckers, Melanerpes formicivorus, a species whose social groups consist of a cooperatively polygynandrous breeding core along with nonbreeding helpers of both sexes that are offspring from prior breeding efforts. Here we examine the life-history differences and the fitness consequences of birds attaining breeder status by either inheriting their natal territory or dispersing to a new territory. Despite significant differences in life history, including the mean territory quality on which individuals bred and mean co-breeder coalition size of breeders, we found no statistical differences in either direct or kin-selected (indirect) fitness benefits for breeders that inherited and dispersed. The incidence of birds engaging in both strategies, inheriting their natal territory and later dispersing, or dispersing but later returning to inherit their natal territory, further reduces the potential direct fitness benefits of inheritance relative to dispersal, since neither precludes the other. Territory inheritance is an important, alternative means of achieving breeding status in this population. However, ecological constraints to dispersal and kin-selected fitness benefits as a helper likely play larger roles driving the acorn woodpecker\u27s extraordinary social system