Pelvic sexual dimorphism and relative neonatal brain size really are related

Primate species in which the neonatal brain size is large relative to the birth canal tend to have more sexually dimorphic pelves: this is a classic comparative relation, discovered by Schultz and Leutenegger. The original work did not correct for phylogenetic nonindependence of the data points; it only partly corrected for body size; it used ratios in both variables, and the size of the female pubis featured in both x- and y-variables. A recent publication by Tague placed a question mark over the validity of the relation. I therefore retested it, correcting for all four statistical defects. A strongly significant statistical relation exists. © 1995 Wiley-Liss, Inc.     

Longevity is associated with relative brain size in birds

Brain size of vertebrates has long been recognized to evolve in close association with basic life‐history traits, including lifespan. According to the cognitive buffer hypothesis, large brains facilitate the construction of behavioral responses against novel socioecological challenges through general cognitive processes, which should reduce mortality and increase lifespan. While the occurrence of brain size–lifespan correlation has been well documented in mammals, much less evidence exists for a robust link between brain size and longevity in birds. The aim of this study was to use phylogenetically controlled comparative approach to test for the relationship between brain size and longevity among 384 avian species from 23 orders. We used maximum lifespan and maximum reproductive lifespan as the measures of longevity and accounted for a set of possible confounding effects, such as allometry, sampling effort, geographic patterns, and life‐history components (clutch size, incubation length, and mode of development). We found that both measures of longevity positively correlated with relative (residual) brain size. We also showed that major diversification of brain size preceded diversification of longevity in avian evolution. In contrast to previous findings, the effect of brain size on longevity was consistent across lineages with different development patterns, although the relatively low strength of this correlation could likely be attributed to the ubiquity of allomaternal care associated with the altricial mode of development. Our study indicates that the positive relationship between brain size and longevity in birds may be more general than previously thought.     

Inter-individual variability contrasts with regional homogeneity in the human brain DNA methylome

The possibility that alterations in DNA methylation are mechanistic drivers of development, aging and susceptibility to disease is widely acknowledged, but evidence remains patchy or inconclusive. Of particular interest in this regard is the brain, where it has been reported that DNA methylation impacts on neuronal activity, learning and memory, drug addiction and neurodegeneration. Until recently, however, little was known about the ‘landscape’ of the human brain methylome. Here we assay 1.9 million CpGs in each of 43 brain samples representing different individuals and brain regions. The cerebellum was a consistent outlier compared to all other regions, and showed over 16 000 differentially methylated regions (DMRs). Unexpectedly, the sequence characteristics of hypo- and hypermethylated domains in cerebellum were distinct. In contrast, very few DMRs distinguished regions of the cortex, limbic system and brain stem. Inter-individual DMRs were readily detectable in these regions. These results lead to the surprising conclusion that, with the exception of cerebellum, DNA methylation patterns are more homogeneous between different brain regions from the same individual, than they are for a single brain region between different individuals. This finding suggests that DNA sequence composition, not developmental status, is the principal determinant of the human brain DNA methylome.     

Relative brain size and ecology in birds

We test the hypothesis that the relative sizes of the different parts of the brain (brain stem, optic lobes, cerebellum and cerebral hemispheres), measured after body size effects have been removed, are associated with differences in behaviour and ecology across bird species.
The results demonstrate that behavioural and ecological correlates of relative brain size are not independent of each other. When the effects of variation in other categories are accounted for, the strongest single effect is due to relatively large brain sizes being associated with altricial development. It is unlikely that this effect is due to the confounding influence of taxonomic associations.
Overall, the results do not provide support for the idea that differences in measures of environmental complexity select for differences in relative brain size.     

The evolution of large brain size in birds is related to social,not genetic,monogamy

There are several hypotheses suggesting that social complexity, including pair bonding, is important in the evolution of increased brain size. I examined whether genetic or social monogamy was related to large brain size in birds. Recent work has indicated that the length and strength of pair bonds are associated with large brain size. I tested several hypotheses for the evolution of large brain size in 42 species of bird by including life history variables in a regression model. A test on 100 phylogenetic trees revealed no phylogenetic signal in brain size. Controlling for body size, a principal components analysis was run on the life history variables and degrees of extra‐pair paternity. The main principal component (PC1) was regressed on brain size revealing a strong, positive association. Social, but not genetic, monogamy was positively related to brain size. Large brain size is related to the selective pressures of procuring extra‐pair copulations whilst maintaining a social partnership. However, other life history variables also loaded positively and significantly on brain size. These results indicate that the evolution of large brain size in birds was driven by several important selective pressures. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 668–678.     

Cerebellum size is related to fear memory and domestication of chickens

Red Junglefowl (Gallus gallus) were selected for divergent levels of fear of humans during eight generations, causing the selection lines to differ in fear levels as well as in the proportional brain and cerebellum masses. Birds from the two lines were then crossed to obtain an F3 intercross in order to study the correlations between brain mass and fear learning. We exposed 105 F3-animals individually to a fear habituation and memory test at 8 days of age, where the reactions to repeated light flashes were assessed on 2 consecutive days. After culling, the absolute and relative sizes of each of four brain regions were measured. Stepwise regression was used to analyse the effects of the size of each brain region on habituation and memory. There were no effects of any brain region on the habituation on day one. However, birds with a larger absolute size of cerebellum had significantly reduced reactions to the fearful stimuli on day two, indicating a better memory of the stimuli. No other regions had significant effects. We conclude that increased cerebellum size may have been important in facilitating chicken domestication, allowing them to adapt to a life with humans.     

Covariation between brain size and immunity in birds: implications for brain size evolution

Parasitism can negatively affect learning and cognition, setting the scene for coevolution between brain and immunity. Greater susceptibility to parasitism by males may impair their cognitive ability, and relatively greater male investment in immunity could compensate for greater susceptibility to parasites, in particular when males have a relatively large brain. We analysed covariation between relative size of immune defence organs and brain in juvenile and adult birds. The relative size of the bursa of Fabricius and the spleen in adults covaried positively with relative brain size across bird species. The relative size of these two immune defence organs covaried with sex differences in relative size of the brain, indicating that the relationship between immune defence and brain size was stronger for males. In contrast, liver and heart size or sexual size dimorphism in size did not covary with immune defence. Thus, species in which males have relatively large brains also have relatively large immune defence organs.     

On the evolution of brain size in relation to migratory behaviour in birds

Migratory birds appear to have relatively smaller brain size compared to sedentary species. It has been hypothesized that initial differences in brain size underlying behavioural flexibility drove the evolution of migratory behaviour; birds with relatively large brains evolved sedentary habits and those with relatively small brains evolved migratory behaviour (migratory precursor hypothesis). Alternative hypotheses suggest that changes in brain size might follow different behavioural strategies and that sedentary species might have evolved larger brains because of differences in selection pressures on brain size in migratory and nonmigratory species. Here we present the first evidence arguing against the migratory precursor hypothesis. We compared relative brain volume of three subspecies of the white-crowned sparrow: sedentary Zonotrichia leucophrys nuttalli and migratory Z. l. gambelii and Z. l. oriantha. Within the five subspecies of the white-crowned sparrow, only Z. l. nuttalli is strictly sedentary. The sedentary behaviour of Z. l. nuttalli is probably a derived trait, because Z. l. nuttalli appears to be the most recent subspecies and because all species ancestral to Zonotrichia as well as all older subspecies of Z. leucophrys are migratory. Compared to migratory Z. l. gambelii and Z. l. oriantha, we found that sedentary Z. l. nuttalli had a significantly larger relative brain volume, suggesting that the larger brain of Z. l. nuttalli evolved after a switch to sedentary behaviour. Thus, in this group, brain size does not appear to be a precursor to the evolution of migratory or sedentary behaviour but rather an evolutionary consequence of a change in migratory strategy.     

Coloniality and migration are related to selection on MHC genes in birds

The major histocompatibility complex (MHC) plays a key role in pathogen recognition as a part of the vertebrate adaptive immune system. The great diversity of MHC genes in natural populations is maintained by different forms of balancing selection and its strength should correlate with the diversity of pathogens to which a population is exposed and the rate of exposure. Despite this prediction, little is known about how life‐history characteristics affect selection at the MHC. Here, we examined whether the strength of balancing selection on MHC class II genes in birds (as measured with nonsynonymous nucleotide substitutions, dN) was related to their social or migratory behavior, two life‐history characteristics correlated with pathogen exposure. Our comparative analysis indicated that the rate of nonsynonymous substitutions was higher in colonial and migratory species than solitary and resident species, suggesting that the strength of balancing selection increases with coloniality and migratory status. These patterns could be attributed to: (1) elevated transmission rates of pathogens in species that breed in dense aggregations, or (2) exposure to a more diverse fauna of pathogens and parasites in migratory species. Our study suggests that differences in social structure and basic ecological traits influence MHC diversity in natural vertebrate populations.     

Evolutionary rates of mitochondrial genomes correspond to diversification rates and to contemporary species richness in birds and reptiles

Rates of biological diversification should ultimately correspond to rates of genome evolution. Recent studies have compared diversification rates with phylogenetic branch lengths, but incomplete phylogenies hamper such analyses for many taxa. Herein, we use pairwise comparisons of confamilial sauropsid (bird and reptile) mitochondrial DNA (mtDNA) genome sequences to estimate substitution rates. These molecular evolutionary rates are considered in light of the age and species richness of each taxonomic family, using a random-walk speciation–extinction process to estimate rates of diversification. We find the molecular clock ticks at disparate rates in different families and at different genes. For example, evolutionary rates are relatively fast in snakes and lizards, intermediate in crocodilians and slow in turtles and birds. There was also rate variation across genes, where non-synonymous substitution rates were fastest at ATP8 and slowest at CO3. Family-by-gene interactions were significant, indicating that local clocks vary substantially among sauropsids. Most importantly, we find evidence that mitochondrial genome evolutionary rates are positively correlated with speciation rates and with contemporary species richness. Nuclear sequences are poorly represented among reptiles, but the correlation between rates of molecular evolution and species diversification also extends to 18 avian nuclear genes we tested. Thus, the nuclear data buttress our mtDNA findings.     

Influence of seasonal changes in relative liver size, condition, relative gonad size and variability in ovarian development in multiple spawning fish species used in environmental monitoring programmes

Basic biological information was collected to assist in understanding the biology of selected small-bodied fishes for use in environmental monitoring programmes. Gonado-somatic index ( I _G) and hepato-somatic index ( I _H) profiles of female blacknose dace Rhinichthys atratulus , mummichog Fundulus heteroclitus and golden shiner Notemigonus crysoleucas were similar, decreasing gradually during the spawning season. I _G and I _H profiles of female northern redbelly dace Phoxinus eos peaked twice suggesting oocyte recruitment was continuous throughout the spawning season. Regressions of ovary mass to adjusted body mass ( M _A) were more variable in dace relative to the other fish species. For female blacknose dace, variability in the relationship between ovary mass and M _A was minimized by selecting either 2 year-old fish and fish that weighed 2–4 g.     

The allometric approach to species differences in brain size

Allometric methods can be used to test quantitative theories of the relationship between brain size and body size across species, and to search for ecological, behavioural, life history, and ontogenetic correlates of brain size. Brain size scales with an allometric exponent of around 0.75 against body size across mammals, but is closer to 0.56 for birds and for reptiles. The slope of the allometric line often varies depending upon the taxonomic level of analysis. However, this phenomenon, at least in mammals, may be a statistical artifact. Brain size for a given body size (relative brain size) varies among orders in birds and mammals, and some dietary associations with relative brain size have been found in particular taxa. Developmental status at birth is the most consistent correlate of relative brain size: precocial neonates have larger brains for a given maternal size than altricial neonates in both birds and mammals. Altricial neonates, however, have more brain growth following birth, and in birds also have larger relative adult brain sizes. Energetic explanations for differences in neonatal brain growth, although attractive on theoretical grounds, have largely failed to stand up to empirical tests.     

Effects of the emerald ash borer invasion on four species of birds

The emerald ash borer (EAB) Agrilus planipennis, first detected in 2002 in the vicinity of Detroit, Michigan, USA, is one of the most recent in a long list of introduced insect pests that have caused serious damage to North American forest trees, in this case ash trees in the genus Fraxinus. We used data from Project FeederWatch, a citizen science program focused on winter bird populations, to quantify the effects of EAB invasion on four species of resident, insectivorous birds known or likely to be EAB predators: three woodpecker species and the white-breasted nuthatch (Sitta canadensis). We compared relative numbers of birds within 50 km of the epicenter of the region where EAB was first detected, an area known to have suffered high ash tree mortality by 2008, to numbers 50–100 km from the epicenter and to control sites within 50 km of five comparable Midwestern cities where damage due to EAB has yet to be severe. We found evidence for significant effects on all four of the species in response to the EAB invasion in the highly impacted region, with red-bellied woodpeckers (Melanerpes carolinus) and white-breasted nuthatches showing numerical increases while downy woodpeckers (Picoides pubescens) and hairy woodpeckers (Picoides villosus) initially declined but exhibited at least temporary increases several years later. Temporal correlation analyses failed to provide support for immigration being a major cause of the elevated numbers in the highly impacted area, and thus these results are consistent with the hypothesis that increases were due to enhanced survival and/or reproduction associated with the EAB invasion within the highly impacted area. Results suggest that the continuing invasion of EAB into new areas is likely to significantly alter avian communities, although not always in ways that will be easy to predict.     

The evolution of hippocampus volume and brain size in relation to food hoarding in birds

Food‐hoarding birds frequently use spatial memory to relocate their caches, thus they may evolve a larger hippocampus in their brain than non‐hoarder species. However, previous studies testing for such interspecific relationships provided conflicting results. In addition, food hoarding may be a cognitively complex task involving elaboration of a variety of brain regions, even outside of the hippocampus. Hence, specialization to food hoarding may also result in the enlargement of the overall brain. In a phylogenetic analysis of distantly related birds, we studied the interspecific association between food hoarding and the size of different brain regions, each reflecting different resolutions. After adjusting for allometric effects, the relative volume of the hippocampus and the relative size of the entire brain were each positively related to the degree of food‐hoarding specialization, even after controlling for migration and brood parasitism. We also found some significant evidence for the relative volume of the telencephalon being associated with food hoarding, but this relationship was dependent on the approach we used. Hence, neural adaptation to food hoarding may favour the evolution of different brain structures.     

Interspecific differences in population trends of Spanish birds are related to habitat and climatic preferences

Aim Animal monitoring programmes have allowed analyses of population trends, most of which now comment on the possible effect of global climate change. However, the relationship between the interspecific variation in population trends and species traits, such as habitat preferences, niche breadth or distribution patterns, has received little attention, in spite of its usefulness in the construction of ecological generalizations. The objectives of this study were: (1) to determine whether there are characteristics shared among species with upwards or downwards trends, and (2) to assess whether population changes agree with what could be expected under global warming (a decrease in species typical of cooler environments). Location The Spanish part of the Iberian Peninsula (c. 500,000 km²) in the south‐western part of the Mediterranean Basin. Methods We modelled recent breeding population changes (1996–2004), in areas without apparent land use changes, for 57 common passerine birds with species‐specific ecological and distributional patterns as explanatory variables. Results One‐half of these species have shown a generalized pattern towards the increase of their populations, while only one‐tenth showed a significant decrease. One half (54%) of the interspecific variability in yearly population trends is explained considering species‐specific traits. Species showing more marked increases preferred wooded habitats, were habitat generalists and occupied warmer and wetter areas, while moderate decreases were found for open country habitats in drier areas. Main conclusions The coherent pattern in population trends we found disagrees with the proposed detrimental effect of global warming on bird populations of western Europe, which is expected to be more intense in bird species inhabiting cooler areas and habitats. Such a pattern suggests that factors other than the increase in temperature may be brought to discussions on global change as relevant components to explain recent changes in biodiversity.     

Sociality, ecology, and relative brain size in lemurs

The social brain hypothesis proposes that haplorhine primates have evolved relatively large brains for their body size primarily as an adaptation for living in complex social groups. Studies that support this hypothesis have shown a strong relationship between relative brain size and group size in these taxa. Recent reports suggest that this pattern is unique to haplorhine primates; many nonprimate taxa do not show a relationship between group size and relative brain size. Rather, pairbonded social monogamy appears to be a better predictor of a large relative brain size in many nonprimate taxa. It has been suggested that haplorhine primates may have expanded the pairbonded relationship beyond simple dyads towards the evolution of complex social groups. We examined the relationship between group size, pairbonding, and relative brain size in a sample of 19 lemurs; strepsirrhine primates that last share a common ancestor with monkeys and apes approximately 75 Ma. First, we evaluated the social brain hypothesis, which predicts that species with larger social groups will have relatively larger brains. Secondly, we tested the pairbonded hypothesis, which predicts that species with a pairbonded social organization will have relatively larger brains than non-pairbonded species. We found no relationship between group size or pairbonding and relative brain size in lemurs. We conducted two further analyses to test for possible relationships between two nonsocial variables, activity pattern and diet, and relative brain size. Both diet and activity pattern are significantly associated with relative brain size in our sample. Specifically, frugivorous species have relatively larger brains than folivorous species, and cathemeral species have relatively larger brains than diurnal, but not nocturnal species. These findings highlight meaningful differences between Malagasy strepsirrhines and haplorhines, and between Malagasy strepsirrhines and nonprimate taxa, regarding the social and ecological factors associated with increases in relative brain size. The results suggest that factors such as foraging complexity and flexibility of activity patterns may have driven selection for increases in brain size in lemurs.     

Bipedal locomotion: effects of speed, size and limb posture in birds and humans

Seven species of ground-dwelling birds (body mass range: 0.045-90 kg) were filmed while walking and running on a treadmill. High-speed light films were also taken of humans to compare kinematic patterns of avian with human bipedalism. Consistent patterns of stride frequency, stride length, step length, duty factor and limb excursion were observed in all species, with most of the variation among species being due to differences in body size. In general, smaller bipeds have higher stride frequencies (α M ^−0.18), shorter stride lengths (α M ^0.38) and more limited ranges of speed within each gait than large bipeds. After normalizing for size (based on Froude number, after Alexander, 1977), remaining kinematic variation is largely due to interspecific differences in posture and relative limb segment lengths. For their size, smaller bipeds have greater step lengths, limb excursion angles and duty factors than large bipeds because of their more crouched posture and greater effective limb length. The most notable differences in limb kinematics between birds and humans occur at the walk-run transition and are maintained as running speed increases. Change of gait is smooth and difficult to discern in birds, but distinct in humans, involving abrupt decreases in step length and duty factor (time of contact) and a corresponding increase in limb swing time. These differences appear to reflect a spring-like run that is stiff in humans (favouring elastic energy recovery) but more compliant in birds (increasing time of ground contact). Differences between birds and humans in balance of the body's centre of mass not only affect femoral orientation and motion, but also affect pattern of limb excursion with speed.