XV.5.5.2 The good genes models can work in the long-term only if the direction of selection pressures affecting the organisms changes in time
The good genes models assume that sexual selection allows females to look for males bearing unsuitable genes and exclude the bearers of these unsuitable genes from reproduction (Fisher 1958). However, these models have a serious drawback. If the selection discriminates against the bearers of a certain trait over a long period of time, it reduces their frequency in the population and thus eliminates polymorphism of this trait from the population. If other factors do not come into play, polymorphism in viability and the degree of expression of (secondary) sexual traits would gradually disappear. As a consequence of the disappearance of polymorphism, the selection pressure on differentiation of males with high and low levels of expression of secondary sexual traits would cease to have any effect on females. Genes for preference for males with a high level of expression of secondary sexual traits would thus cease to be selected and would gradually disappear from the population. This principle is especially intensely applied in species in which females select males or, to be more exact, their genes, as they get nothing more from them, together on courting grounds (leks). In a great many species, the females copulate here, one after another, with a single, the best male and the other males have no role in reproduction. If the fitness of males were to have non-zero heritability, all the variability in the relevant genes would be eliminated in a very few generations and the females would have no reason for discriminating between males. Then females that did not lose time and energy in selecting males at a lek would be at an advantage, so that the relevant female genes for selectivity and the male genes for the secondary sexual traits would rapidly disappear from the population. The long-term persistence of a particular reproduction system and variability in genes for secondary sexual traits is called the lek paradox. A number of more or less probable mechanisms have been gradually proposed to resolve this paradox (Randerson, Jiggins, & Hurst 2000; Williams 1992){13754}.
It is apparent that the good genes model can work only if the object of selection consists in those traits whose selection value continues to change. These traits could include, for example, resistance to parasites. This can even exhibit negative heritability, i.e. genotypes that are advantageous in one generation automatically become disadvantageous in the following generation (see XIII.3.2.2.3). The winter habitat hypothesis assumes that the selection value for genes controlling the site of the winter habitat and the route of the spring and autumn migration of birds change through changes in the climate and weather in various parts of the world. Similarly, the selection value for genes controlling some patterns of social behavior can change in dependence on changes in abiotic factors and on the behavior of the other individuals in the population.