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It may be that lower-anity antibodies pri- marily interfere with attachment to host cells by covering most viral attachment sites buy 100mg clomid fast delivery menopause 6 months without a period. By contrast purchase clomid line women's health june 2012, higher-anity antibodies may interfere primarily with fusion and entry to host cells buy clomid with a visa womens health 40 is the new 20, and such steric interference at the cell surface requires a lower density ofbound antibody generic penegra 100 mg. When virions attachtocellsurfaces order 160mg super avana mastercard,the lower-anity epitopes may lose alargerfractionofbound antibody than higher-anity epitopes. Synergism occurs when simultaneous binding by two antibodies causes higher neutralization than expected by adding the eects of each anti- body when bound alone. Thus, the tness eect of an amino acid sub- stitution may depend both on the reduced anity fortheconforming antibody and on the context of other antibody-epitope combinations for that pathogen genotype. Structural studies locate particular amino acid sites in their three-dimensional context. Experimental evolution substitutes amino acids in response to immune pressure, altered cellular receptors, in- terference with the viral receptor binding site, or changed kinetics that arise in cell culture. Binding anity and kinetics ofneutralization relate amino acid substitutions to components of tness. In this section, I briey list a few additional studies of experimental evolution. Experimental deletion of the B cell response led to an absence of aminoacidsubstitutions in the presumed antibody epitopes, demonstrating that substitutions in surface proteins arise in response to antibodies rather than cell tropism. Not surprisingly, escape mutants do arise frequently with amino acid substitutions in the immunodominant surface antigens (Gow and Mutimer 2000). Antigenic change in response to antibody pressure can change polymerase function, and substitutions in the polymerase in re- sponse to nucleoside analog drugs canchange antigenic properties of surface proteins. The mapping of amino acidsubstitutions to tness may be rather complex in this case. Amino acid substitutions in measles hem- agglutinin appear to alter both antigenicity and neurovirulence. Measles virus also appears to change its binding anity for dierent cellularreceptors during adaptation to cell culture (Nielsen et al. The amino acid changes associated with receptor anity occur in the surface hemagglutinin protein. Thelife cycle of arthropod-borne viruses (arboviruses) typically al- ternates between vertebrate hosts and blood-feeding arthropod vectors. However, many stud- ies have reported a high degree of antigenic conservation and slow rates of molecular evolution (reviewed byCooper and Scott 2001). Cooper and Scott (2001) used experimental evolution to study how alternating hosts potentially constrain adaptive change.

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It is not so easy to imagine the spatio- temporal landscape because the spread of each variant has diering quantitative eects on the dynamics ofothervariants purchase clomid us breast cancer wigs. One simple anal- ogy with age structure hints at thesortofprocesses that may occur buy 25mg clomid with mastercard menstruation tiredness. In inuenza buy discount clomid on line menstrual volume, it may be that children have immunodominant memory focused on only one or a few antigenic sites generic 50mg clomid with mastercard. In the simplied example Idiscussed above order cheapest cialis jelly and cialis jelly, at rst a virus strain with two sites, A/B,spreads. Some children develop immunodominant memory against A;otherchil- dren develop immunodominant memory against B. The key is that dierent classes of hosts provide a pathway of connectivity by which single mutations of the virus can eventually spread through the entire population. This spatial distribution of immunological memory creates astepwisepathway of connectivity for a parasite. For example, if a parasite A /B rst invades patch 1, then it can by a single mutation change into A/B and attack patch 2. First, patches 1 and 3 uctuate between A/B and A /B,whereaspatches 2and4uctuate between A /B and A/B. Thesepatchidentities occur because immunological memory to both antigens imposes a barrier to any variant except the type with changes at both sites. With spatial structure, alternating regions of the host population can be dominated by the dierent pairwise sets of parasite strains. I will return to these issues in the next chapter, which focuses on the population structure of antigenically variable parasites. Mathematical models could be developed to explore the interactions between antigenic variation and spatiotemporal dynamics. Some spatial data exist for inuenza, but the scale of sampling and the measurement of cross-reactivity prob- ably need to be enhanced before much can be concluded. But eventually methods will improve for typing strains, and more data will become available. Rather than repeatallofthose issues, I list here some hypotheses about kinetics that deserve study both empirically and math- ematically. Many patterns of antigenic variation turn on these rate pro- cesses that drive evolutionary dynamics. Consider two species, a long-lived species with an average life span of L years, and a short-lived species with an average life span of S years. By contrast, if S = 7, then newborns replace approximately 1/7th of the population each year. Immunological memory decays faster at the pop- ulation level in short-lived than in long-lived species, perhaps reducing the relativetnessadvantage of antigenic variantsinshort-livedcom- pared with long-lived hosts.

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