Reproductive sharing in animal societies: reproductive incentives or incomplete control by dominant breeders?

Optimal skew models explain reproductive sharing within social groups as resulting from reproductive incentives given by controlling dominants to subordinates in return for peaceful cooperation. We explore two versions of an alternative, the incomplete control model, for the evolution of reproductive sharing within groups. In this model, dominants have only limited control over the allocation of reproduction and must expend effort to increase their share of the total group output We show that, when the relatedness between dominant and subordinate is symmetrical, (1) the subordinate's fraction of reproduction either increases with, or is insensitive to, the subordinate's genetic relatedness, r, to the dominant in both versions of the incomplete control model, whereas the subordinate's fraction of reproduction decreases with increasing r in the optimal skew model, (2) the subordinate's share of reproduction in the incomplete control model must exceed that in the optimal skew model, and (3) ecological factors affecting solitary breeding success do not directly affect the subordinate's share of reproduction in incomplete control model but do in the optimal skew model. When dominant-subordinate relatedness is asymmetrical (as is often the case in parent-offspring associations), the incomplete control model predicts no reproduction by the subordinate offspring regardless of group size for groups containing any mixture of unrelated and full-sibling subordinates, whereas the optimal skew models predict that such reproduction is possible when the group size is three or more. The available evidence indicates a negative relationship between relatedness and a subordinate's reproductive share in both vertebrate and hymenopteran societies, apparently supporting the predictions of the optimal skew, not incomplete control, class of models. However, such a negative relationship is not necessarily inconsistent with the incomplete control model when, as is true for some vertebrate studies, it results from a comparison of skews in genetically monogamous, nonincestuous groups of parents and their offspring (asymmetric relatednesses) with skews in groups of nonkin (symmetric relatednesses). Both models predict higher skews in parent-offspring associations. Occasional reproduction by subordinate offspring in groups of asymmetrical relatedness when such groups are larger than dyads is more consistent with the optimal skew model, however. Overall, current data on reproductive skew and its relationships to intragroup aggression and ecological constraints support the optimal skew model, but more data are needed to rule out the incomplete control model. These models are examples of two different general views of intrasocietal evolution: the tug-of-war view, in which group members engage in a struggle over resources, and the transactional view, in which group members exchange parcels of reproduction to induce beneficial behavior from each othe

γ-Glutamyltransferase, but not markers of hepatic fibrosis, is associated with cardiovascular disease in older people with type 2 diabetes mellitus: the Edinburgh Type 2 Diabetes Study

AIMS/HYPOTHESIS: We examined the association of prevalent and incident cardiovascular disease (CVD) with chronic liver disease in a cohort of community-based people with type 2 diabetes, in order to clarify the relationship between these two important conditions. METHODS: 1,066 participants with type 2 diabetes aged 60–75 years underwent assessment of a range of liver injury markers (non-specific injury, steatosis, steatohepatitis, fibrosis, portal hypertension). Individuals were followed up for incident cardiovascular events. RESULTS: At baseline there were 370/1,033 patients with prevalent CVD, including 317/1,033 with coronary artery disease (CAD). After a mean follow-up of 4.4 years there were 44/663 incident CVD events, including 27/663 CAD events. There were 30/82 CVD-related deaths. Risk of dying from or developing CVD was no higher in participants with steatosis than in those without (HR 0.90; 95% CI 0.40, 2.00; p > 0.05). The only notable relationship was with γ-glutamyltransferase (GGT) (incident CVD: adjusted HR for doubling GGT 1.24 [95% CI 0.97, 1.59] p = 0.086; incident CAD: adjusted HR 1.33 [95% CI 1.00, 1.78] p = 0.053), suggesting that in our study population, chronic liver disease may have little effect on the development of, or mortality from, CVD. CONCLUSIONS/INTERPRETATION: An independent association between GGT and CVD warrants further exploration as a potentially useful addition to current cardiovascular risk prediction models in diabetes. However, overall findings failed to suggest that there is a clinical or pathophysiological association between chronic liver disease and CVD in elderly people with type 2 diabetes. ELECTRONIC SUPPLEMENTARY MATERIAL: The online version of this article (doi:10.1007/s00125-015-3575-y) contains peer-reviewed but unedited supplementary material, which is available to authorised users