Despite the wealth of empirical and theoretical studies, the origin and maintenance of cooperation is still an evolutionary riddle. In this context, ecological life-history traits which affect the efficiency of selection may play a role despite being often ignored. We consider here species such as bacteria, fungi, invertebrates and plants which exhibit resting stages in the form of a quiescent state or a seed bank. When quiescent, individuals are inactive and reproduce upon activation, while under seed bank parents produce offspring remaining dormant for different amount of time. We assume weak frequency-dependent selection modeled using game-theory and the prisoner's dilemma (cooperation/defect) as payoff matrix. The cooperators and defectors are allowed to evolve different quiescence or dormancy times. By means of singular perturbation theory we reduce the model to a one-dimensional equation resembling the well known replicator equation, in which the gain functions are scaled with lumped parameters reflecting the time scale of the resting state of the cooperators and defectors. If both time scales are identical cooperation cannot persist in a homogeneous population. If, however, the time scale of the cooperator is distinctively different from that of the defector, cooperation may become a locally asymptotically stable strategy. Interestingly enough, in the seed bank case the cooperator needs to become active faster than the defector, while in the quiescent case the cooperator has to be slower. We use adaptive dynamics to identify situations where cooperation may evolve and form a convergent stable ESS. We conclude by highlighting the relevance of these results for many non-model species and the maintenance of cooperation in microbial, invertebrate or plant populations.

中文翻译：

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更好的合作者是处于休眠状态还是处于静止状态？

尽管有大量的经验和理论研究，合作的起源和维持仍然是一个进化之谜。在这种情况下，尽管经常被忽视，但影响选择效率的生态生活史特征却可能发挥作用。我们在这里考虑以静止状态或种子库的形式表现出静止阶段的物种，例如细菌，真菌，无脊椎动物和植物。处于静止状态时，个体处于非活动状态，并在激活后繁殖，而在种子库之下，父母则产生后代保持休眠状态的时间不同。我们假设使用基于博弈论和囚徒困境（合作/缺陷）作为回报矩阵的弱频率依赖选择模型。允许合作者和叛逃者发展不同的静止或休眠时间。借助奇异摄动理论，我们将模型简化为类似于众所周知的复制子方程的一维方程，其中增益函数通过集总参数缩放，该集总参数反映了合作者和叛逃者的静止状态的时间尺度。如果两个时间尺度都相同，那么合作就不可能在同质人群中持续进行。但是，如果合作者的时间尺度与叛逃者的时间尺度明显不同，则合作可能会成为局部渐近稳定的策略。有趣的是，在种子库的情况下，合作者需要比叛逃者更快地活跃起来，而在静态的情况下，合作者必须更慢。我们使用自适应动力学来确定合作可能演变并形成收敛稳定的ESS的情况。