SELECTION ON MENOPAUSE IN TWO PREMODERN HUMAN POPULATIONS: NO EVIDENCE FOR THE MOTHER HYPOTHESIS

Evolutionary theory suggests that natural selection should synchronize senescence of reproductive and somatic systems. In some species, females show dramatic discordance in senescence rates in these systems, leading to a clear menopause coupled with prolonged postreproductive life span. The Mother Hypothesis proposes that menopause evolved to avoid higher reproductive‐mediated mortality risk in late‐life and ensure the survival of existing offspring. Despite substantial theoretical interest, the critical predictions of this hypothesis have never been fully tested in populations with natural fertility and mortality. Here, we provide an extensive test, investigating both short‐ and long‐term consequences of mother loss for offspring, using multigenerational demographic datasets of premodern Finns and Canadians. We found no support for the Mother Hypothesis. First, although the risk of maternal death from childbirth increased from middle age, the risk only reached 1–2% at age 50 and was predicted to range between 2% and 8% by 70. Second, offspring were adversely affected by maternal loss only in their first two years (i.e., preweaning), having reduced survival probability in early childhood as well as ultimate life span and fitness. Dependent offspring were not affected by maternal death following weaning, either in the short‐ or long‐term. We suggest that although mothers are required to ensure offspring survival preweaning in humans, maternal loss thereafter can be compensated by other family members. Our results indicate that maternal effects on dependent offspring are unlikely to explain the maintenance of menopause or prolonged postreproductive life span in women.     

Evolutionary demography of agricultural expansion in preindustrial northern Finland

A shift from nomadic foraging to sedentary agriculture was a major turning point in human evolutionary history, increasing our population size and eventually leading to the development of modern societies. We however lack understanding of the changes in life histories that contributed to the increased population growth rate of agriculturalists, because comparable individual-based reproductive records of sympatric populations of agriculturalists and foragers are rarely found. Here, we compared key life-history traits and population growth rate using comprehensive data from the seventieth to nineteenth century Northern Finland: indigenous Sami were nomadic hunter-fishers and reindeer herders, whereas sympatric agricultural Finns relied predominantly on animal husbandry. We found that agriculture-based families had higher lifetime fecundity, faster birth spacing and lower maternal mortality. Furthermore, agricultural Finns had 6.2% higher annual population growth rate than traditional Sami, which was accounted by differences between the subsistence modes in age-specific fecundity but not in mortality. Our results provide, to our knowledge, the most detailed demonstration yet of the demographic changes and evolutionary benefits that resulted from agricultural revolution.     

Generation of comprehensive transposon insertion mutant library for the model archaeon, <Emphasis Type="Italic">Haloferax volcanii</Emphasis>, and its use for gene discovery

Background

Archaea share fundamental properties with bacteria and eukaryotes. Yet, they also possess unique attributes, which largely remain poorly characterized. Haloferax volcanii is an aerobic, moderately halophilic archaeon that can be grown in defined media. It serves as an excellent archaeal model organism to study the molecular mechanisms of biological processes and cellular responses to changes in the environment. Studies on haloarchaea have been impeded by the lack of efficient genetic screens that would facilitate the identification of protein functions and respective metabolic pathways.

Results

Here, we devised an insertion mutagenesis strategy that combined Mu in vitro DNA transposition and homologous-recombination-based gene targeting in H. volcanii. We generated an insertion mutant library, in which the clones contained a single genomic insertion. From the library, we isolated pigmentation-defective and auxotrophic mutants, and the respective insertions pinpointed a number of genes previously known to be involved in carotenoid and amino acid biosynthesis pathways, thus validating the performance of the methodologies used. We also identified mutants that had a transposon insertion in a gene encoding a protein of unknown or putative function, demonstrating that novel roles for non-annotated genes could be assigned.

Conclusions

We have generated, for the first time, a random genomic insertion mutant library for a halophilic archaeon and used it for efficient gene discovery. The library will facilitate the identification of non-essential genes behind any specific biochemical pathway. It represents a significant step towards achieving a more complete understanding of the unique characteristics of halophilic archaea.

     

Rapid recovery of invertebrate communities after ecological restoration of boreal mires

Mire degradation due to drainage for forestry results in the loss of mire specialist species. To halt the loss in biodiversity, ecological restoration is needed and already implemented. However, a major challenge in ecological restoration is whether actions taken have the desired outcome. Key abiotic and biotic conditions for the successful restoration of invertebrate communities can be identified by testing the “Field of Dreams” hypothesis, which postulates that if a habitat is successfully restored, species will return. This study was conducted in nine boreal mires located in Eastern Finland, 1–3 years after restoration. Parts of each mire were drained for forestry during the 1960s and 1970s, and restored in 2003–2006. Two 250 m transects were established in each of three treatments (pristine, drained, restored) per mire. We used pitfall trapping to sample carabid beetles and spiders, sweep netting to sample micromoths and crane flies and counts along the transects to sample macromoths. Vegetation cover, water table level, and climatic variables were measured along all transects. Mire specialist species and invertebrate communities responded positively to restoration and negatively to drainage, whereas generalists showed varied responses. In addition, mire specialists were associated with high cover of Sphagnum mosses and with low numbers of tall trees (>3 m). Therefore, to successfully restore populations and communities of mire specialist invertebrates, maintaining environmental conditions that favor the growth of Sphagnum mosses, rewetting the sites and removing larger trees are necessary measures.     

Will granny save me? Birth status,survival, and the role of grandmothers in historical Finland

Grandmothers play a crucial role in families enhancing grandchild wellbeing and survival but their effects can be context-dependent, and the children born in poor conditions are most likely to benefit from the investments made by helping grandmothers. In this study, we examined, for the first time, whether grandmothers' presence modified associations between adverse birth status and survival up to 5 years of age. In detail, we verified, whether (i) firstborns, (ii) twins, (iii) children born within 24 months after their sibling, and (iv) children followed by short interval (i.e. their younger sibling was born within 24 months) survived better when either their maternal, paternal, or both grandmothers were present. Moreover, we evaluated whether illegitimate children survived better when the maternal grandmother was present. We used an extensive and largely pre-industrial demographic dataset collected from parish population registers kept by the Lutheran Church of Finland from years 1730–1895. We show that although grandmother presence cannot mitigate adverse effects of many poorer birth conditions, grandchildren whose next sibling was born after a short interval survived better when the maternal grandmother was present. Taken together, these findings highlight an important role of grandmothers in compensating the mother's investment in the new baby, thus enabling overall faster successful reproductive rate of mothers. Whilst the opportunity for grandmothers to mitigate the risks of adverse birth statuses is limited, this study does show - through the beneficial effect on survival for those with a short subsequent birth interval - that grandmothers can increase their daughters' and their own reproductive success.     

Masculinization of the eruption pattern of permanent mandibular canines in opposite sex twin girls

The aim of this study is to explore the effect of prenatal androgenization on the clinical eruption of permanent teeth expressing dimorphism and bimaturism. The eruption curves of permanent teeth (except third molars), including those that make up the canine complex (permanent canines, lower first premolars), are compared among opposite sex twins (OS twins) relative to single‐born boys and girls. The comparisons are made with regard to three phases of eruption (pierced mucosa, half‐ erupted, and completely erupted) from a cross‐sectional sample of dental casts, using Kaplan–Meier survival and Cox regression analyzes. The casts were collected from 2159 school children from the US Collaborative Perinatal Project, including 39 pairs of OS‐twins, of which 12 pairs (30.8%) were Euro‐Americans and 27 pairs (69.2%) were of African‐American ancestry. The eruption patterns of the incisors, upper first molars, and lower canines were found to be significantly masculinized (delayed) among OS twin girls. The differences in most other teeth were either not significant, or the number of observations of active eruption phases were too few, such as in the upper first molars and incisors, to yield strong evidence and meaningful results. The masculinization of the tooth eruption pattern in OS twin girls is intriguing because of the lower canine responses during puberty, as well as canine primordial formation during early fetal androgenization of their co‐twin during the 8th to 14th gestational weeks. The present results offer a challenge for future research exploring tooth eruption mechanisms, and may also highlight some cases of delayed or ectopic canines, which are biased toward females. Am J Phys Anthropol 151:566–572, 2013. © 2013 Wiley Periodicals, Inc.     

Maternal risk of breeding failure remained low throughout the demographic transitions in fertility and age at first reproduction in Finland

Radical declines in fertility and postponement of first reproduction during the recent human demographic transitions have posed a challenge to interpreting human behaviour in evolutionary terms. This challenge has stemmed from insufficient evolutionary insight into individual reproductive decision-making and the rarity of datasets recording individual long-term reproductive success throughout the transitions. We use such data from about 2,000 Finnish mothers (first births: 1880s to 1970s) to show that changes in the maternal risk of breeding failure (no offspring raised to adulthood) underlay shifts in both fertility and first reproduction. With steady improvements in offspring survival, the expected fertility required to satisfy a low risk of breeding failure became lower and observed maternal fertility subsequently declined through an earlier age at last reproduction. Postponement of the age at first reproduction began when this risk approximated zero-even for mothers starting reproduction late. Interestingly, despite vastly differing fertility rates at different stages of the transitions, the number of offspring successfully raised to breeding per mother remained relatively constant over the period. Our results stress the importance of assessing the long-term success of reproductive strategies by including measures of offspring quality and suggest that avoidance of breeding failure may explain several key features of recent life-history shifts in industrialized societies.     

Metabolism of Vertebrate Amino Sugars with N-Glycolyl Groups: RESISTANCE OF α2-8-LINKED N-GLYCOLYLNEURAMINIC ACID TO ENZYMATIC CLEAVAGE

The sialic acid (Sia) N-acetylneuraminic acid (Neu5Ac) and its hydroxylated derivative N-glycolylneuraminic acid (Neu5Gc) differ by one oxygen atom. CMP-Neu5Gc is synthesized from CMP-Neu5Ac, with Neu5Gc representing a highly variable fraction of total Sias in various tissues and among different species. The exception may be the brain, where Neu5Ac is abundant and Neu5Gc is reported to be rare. Here, we confirm this unusual pattern and its evolutionary conservation in additional samples from various species, concluding that brain Neu5Gc expression has been maintained at extremely low levels over hundreds of millions of years of vertebrate evolution. Most explanations for this pattern do not require maintaining neural Neu5Gc at such low levels. We hypothesized that resistance of α2-8-linked Neu5Gc to vertebrate sialidases is the detrimental effect requiring the relative absence of Neu5Gc from brain. This linkage is prominent in polysialic acid (polySia), a molecule with critical roles in vertebrate neural development. We show that Neu5Gc is incorporated into neural polySia and does not cause in vitro toxicity. Synthetic polymers of Neu5Ac and Neu5Gc showed that mammalian and bacterial sialidases are much less able to hydrolyze α2-8-linked Neu5Gc at the nonreducing terminus. Notably, this difference was not seen with acid-catalyzed hydrolysis of polySias. Molecular dynamics modeling indicates that differences in the three-dimensional conformation of terminal saccharides may partly explain reduced enzymatic activity. In keeping with this, polymers of N-propionylneuraminic acid are sensitive to sialidases. Resistance of Neu5Gc-containing polySia to sialidases provides a potential explanation for the rarity of Neu5Gc in the vertebrate brain.     

Elder brothers affect the life history of younger siblings in preindustrial humans: social consequence or biological cost?

Sex-specific sibling interactions are potentially important in human ecology. It is well established that in patrilineal societies that sons suffer from the presence of brothers because of competition for inheritance. However, offspring (of both sexes) might also suffer from being born after an elder brother because of the greater costs sons may entail for their mother. Evidence that the cost of producing sons is higher has been gained from studies of ungulates and humans, with some of this cost being manifested as lower birthweight or reproductive performance of offspring born following a male. Using church record data from preindustrial Finland, we shed light on this process by investigating the demographic ‘mechanisms' by which offspring born following an elder brother are compromised. First, we show that, for both men and women in this population, being born after an elder male sibling is associated with reduced probability of reproducing, a later age at first reproduction, and longer interbirth intervals. Second, we show that the primary effect of interest is a reduced probability of reproducing in those born after an elder brother (even among only those who married). Finally, we show that the total number of elder brothers who survived to adulthood has a negative effect on male offspring only, and this effect is independent of the elder brother effect above. We highlight that differences in the success of human offspring are not always social in origin as is often perceived but can also be biological, resulting from differential costs for mothers of producing male versus female offspring.