In most organisms, genome haploidization requires reciprocal DNA exchanges (crossovers) between replicated parental homologs to form bivalent chromosomes. These are resolved to their four constituent chromatids during two meiotic divisions. In female mammals, bivalents are formed during fetal life and remain intact until shortly before ovulation. Extending this period beyond 35 years greatly increases the risk of aneuploidy in human oocytes, resulting in a dramatic increase in infertility, miscarriage, and birth defects, most notably trisomy 21. Bivalent chromosomes are stabilized by cohesion between sister chromatids, which is mediated by the cohesin complex. In mouse oocytes, cohesin becomes depleted from chromosomes during female aging. Consistent with this, premature loss of centromeric cohesion is a major source of aneuploidy in oocytes from older women. Here, we propose a mechanistic framework to reconcile data from genetic studies on human trisomy and oocytes with recent advances in our understanding of the molecular mechanisms of chromosome segregation during meiosis in model organisms.

Immune defenses are expected to be crucial for survival under the considerable parasite pressures experienced by wild animals. However, our understanding of the association between immunity and fitness in nature remains limited due to both the complexity of the vertebrate immune system and the often‐limited availability of immune reagents in nonmodel organisms. Here, we use methods and reagents developed by veterinary researchers for domestic ungulates on blood samples collected from a wild Soay sheep population, to evaluate an unusually broad panel of immune parameters. Our evaluation included different innate and acquired immune cell types as well as nematode parasite‐specific antibodies of different isotypes. We test how these markers correlate with one another, how they vary with age‐group and sex, and, crucially, whether they predict overwinter survival either within or among demographic groups. We found anticipated patterns of variation in markers with age, associated with immune development, and once these age trends were accounted for, correlations among our 11 immune markers were generally weak. We found that females had higher proportions of naïve T cells and gamma–delta T cells than males, independent of age, while our other markers did not differ between sexes. Only one of our 11 markers predicted overwinter survival: sheep with higher plasma levels of anti‐nematode IgG antibodies were significantly more likely to survive the subsequent high mortality winter, independent of age, sex, or weight. This supports a previous finding from this study system using a different set of samples and shows that circulating antibody levels against ecologically relevant parasites in natural systems represent an important parameter of immune function and may be under strong natural selection. Our data provide rare insights into patterns of variation among age‐ and sex groups in different T‐cell subsets and antibody levels in the wild, and suggest that certain types of immune response—notably those likely to be repeatable within individuals and linked to resistance to ecologically relevant parasites—may be most informative for research into the links between immunity and fitness under natural conditions.

We consider a stochastic model for evolution of group-structured populations in which interactions between group members correspond to the Prisoner's Dilemma or the Hawk-Dove game. Selection operates at two organization levels: individuals compete with peer group members based on individual payoff, while groups also compete with other groups based on average payoff of group members. In the Prisoner's Dilemma, this creates a tension between the two levels of selection, as defectors are favored at the individual level, whereas groups with at least some cooperators outperform groups of defectors at the between-group level. In the limit of infinite group size and infinite number of groups, we derive a non-local PDE that describes the probability distribution of group compositions in the population. For special families of payoff matrices, we characterize the long-time behavior of solutions of our equation, finding a threshold intensity of between-group selection required to sustain density steady-states and the survival of cooperation. When all-cooperator groups are most fit, the average and most abundant group compositions at steady-state range from featuring all-defector groups when individual-level selection dominates to featuring all-cooperator groups when group-level selection dominates. When the most fit groups have a mix of cooperators and defectors, then the average and most abundant group compositions always feature a smaller fraction of cooperators than required for the optimal mix, even in the limit where group-level selection is infinitely stronger than individual-level selection. In such cases, the conflict between the two levels of selection cannot be decoupled, and cooperation cannot be sustained at all in the case where between-group competition favors an even mix of cooperators and defectors.