This study establishes a theoretical framework for differentiating among possible behavioral mechanisms whereby sexual communication of moths is disrupted in crops treated with point sources of pheromone. The major mechanisms recognized in the mating disruption literature fall into two main categories: competitive (competitive attraction = false-plume-following) and non-competitive (camouflage, desensitization, and sensory imbalance). Each disruption mechanism has been precisely defined verbally, and then the distinguishing characteristics of the two categories were defined mathematically. The sets of predictions associated with each category were visualized by graphical plots of mathematical simulations. Profiles of simulated male visitation rates to pheromone-baited traps deployed in pheromone-treated crops were graphed against density of pheromone dispensers by using various types of axes. Key traits of non-competitive attraction are as follows: concave profiles on untransformed axes, with an asymptotic approach to zero catch of male moths in traps; a straight line with positive slope when 1/catch is plotted against dispenser density (Miller-Gut plot); and a straight line with negative slope when catch is plotted against dispenser density * catch (Miller-de Lame plot). Key traits of non-competitive disruption profiles include: an initial linear disruption profile on untransformed axes; a concave Miller-Gut plot; and a recurving Miller-de Lame plot. These differences in profiles provide a basis for distinguishing competitive from non-competitive mechanisms when analyzing disruption profiles from field experiments. Slopes and intercepts of these secondary plots can also reveal both male and female moth densities, if the relative attractiveness of traps, females, and dispensers is known. The absolute value of the slope of the Miller-de Lame plot is a measure of each dispenser's activity (D(a)) for suppressing catch of male moths in traps. An application activity (D(Aa)) for a given dispenser can be calculated by multiplying D(a) by the number of such dispensers applied per hectare of crop.

Eleven out of 13 disruption profiles (plots of dispenser density vs. male catch) from moth sex pheromone literature were consistent with a competitive-attraction mechanism, in which dispensers attract males and thereby divert them from females. Mean dispenser activity (D(a)) across all competitive-attraction cases was 0.04 +/- 0.06 (SD); values ranged from 0.0005 for a tiny laminated flake dispenser of racemic disparlure targeting gypsy moth to 0.2 for polyethylene tube dispensers used against lightbrown apple moth. A dispenser application activity (D(Aa)) can be calculated by multiplying D(a) by the number of such dispensers applied per hectare of crop. The highest dispenser application activity (D(Aa)) values approached 200 and corresponded to >99% inhibition of catches of male moths in monitoring traps. Relative to the D(Aa) scale, % inhibition of catches of male moths compressed and obscured large differences in D(Aa) when % disruption exceeded 90%. For cases of competitive attraction, these two efficacy scales can be interconverted by using the formula: D(Aa) approximately = 100/(100 minus % disruption). When disruptive point sources of pheromone were directly observed, male moths were seen approaching pheromone dispensers whose disruption profiles matched competitive attraction. Two cases fit non-competitive disruption mechanisms, which include camouflage, desensitization (adaptation and/or habituation), and sensory imbalance. In these cases, pheromone was released at rates higher than for cases of disruption by competitive attraction. Practical ramifications of the finding that competitive attraction appears to be the prevalent mechanism for moth mating disruption by pheromone point sources are listed. We believe that the congruence of diverse sets of mating disruption field data with explicit a priori predictions validates competitive-attraction theory. The analytical tools and principles governing competitive attraction that were uncovered during this study of mating disruption of moths should be generally applicable to competitive-attraction phenomena.

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