Spite is the most surprising prediction of inclusive fitness theory because it suggests that a gene can be favored by natural selection despite causing harm to both the individuals that carry it and those around them. A gene for spite can only be favored because of negative relatedness, which means that the actor that carries the gene is less likely to share the gene for spite with the surrounding recipients than the random expectation. While positive relatedness can be simply reduced to the intuitive concept of kinship, negative relatedness is deeply counterintuitive. Here I clarify that negative relatedness is frequency dependent, and I identify a hidden assumption in its widely used formula. Accordingly, while the well-studied “lighter” side of inclusive fitness (with helping behaviors and positive relatedness) is dominated by traits that are favored under kin selection, I predict that the understudied “darker” side of inclusive fitness (with harming behaviors and negative relatedness) is dominated by traits that are favored under greenbeard/kind selection—and I discuss the existing evidence that tentatively supports this hypothesis.

中文翻译：

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恶意和负面关联的几何

尽善尽美是包容性适应性理论的最令人惊讶的预测，因为它表明，尽管对携带该基因的个人和周围的人们都造成了伤害，但该基因仍可被自然选择所青睐。仅出于消极相关性而偏爱一个基因，这意味着携带该基因的演员与周围的接受者相比，与随机期望相比，不太可能与该对象共享该基因。虽然正面关系可以简单地简化为亲属关系的直觉概念，但负面关系却与直觉相反。在这里，我澄清了负相关性与频率有关，并且在其广泛使用的公式中确定了一个隐藏的假设。因此，