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When the expected drift time of the intermediate genotype is short, we can neglect the changing background fitness due to the uphill mutant during this drift time.
However, when s u is small or δ i is large, this background fitness changes slowly compared to the intermediate drift time.
These interference effects are possible because of an extensive genetic linkage across influenza's HA (Strelkowa and Lässig, 2012) and arise from variation in the background fitness of viral strains and from variation in the fitness effects of subsequent mutations.
The much longer predicted persistence times of our lethal transgenes under permissive conditions suggests that it is possible to design constructs and generate transgene insertion sites where such background fitness costs can be minimized.
As such, even though the genetic change is one and the same, the change in binding avidity that results can be thought of as increasing the background fitness of a virus strain, whereas the change in antigenicity that results can be considered as in this paper.
Whether glycosylation sites accumulate over evolutionary time (as in the case of H3N2 in humans [ Blackburne et al., 2008]) or do not (as in the case of H1N1 in humans [ Das et al., 2011]) therefore will depend on how these sites impact the background fitness of virus strains.
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Alternatively, there is a simple and relevant parameter regime in which background fitnesses change slowly.
Of course, it's a bit dangerous to start recommending training to people where you can't even see them, you have no idea of their background or fitness.
Ms Thomson said: "The issues fall into three categories which are firstly around the driver of the vehicle, his medical background, his fitness to hold the licence, his employment record and training.
His background in fitness helps, too.
Andrews, who has a master's degree in psychology and a background in fitness, runs the Institute of Sports Performance in Houston.
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