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Introduction: Songbirds, especially corvids, and parrots are remarkably intelligent. Their cognitive skills are on par with primates and their brains contain primate-like numbers of neurons concentrated in high densities in the telencephalon. Much less is known about cognition and neuron counts in more basal bird lineages. Here, we focus on brain cellular composition of galliform birds, which have small brains relative to body size and a proportionally small telencephalon and are often perceived as cognitively inferior to most other birds. METHODS: We use the isotropic fractionator to assess quantitatively the numbers and distributions of neurons and nonneuronal cells in 15 species of galliform birds and compare their cellular scaling rules with those of songbirds, parrots, marsupials, insectivores, rodents, and primates. RESULTS: On average, the brains of galliforms contain about half the number of neurons found in parrot and songbird brains of the same mass. Moreover, in contrast to these birds, galliforms resemble mammals in having small telencephalic and dominant cerebellar neuronal fractions. Consequently, galliforms have much smaller absolute numbers of neurons in their forebrains than equivalently sized songbirds and parrots, which may limit their cognitive abilities. However, galliforms have similar neuronal densities and neuron counts in the brain and forebrain as equally sized non-primate mammals. Therefore, it is not surprising that cognitive abilities of galliforms are on par with non-primate mammals in many domains. CONCLUSION: Comparisons performed in this study demonstrate that birds representing distantly related clades markedly differ in neuronal densities, neuron numbers, and the allocation of brain neurons to major brain divisions. In analogy with the concept of volumetric composition of the brain, known as the cerebrotype, we conclude that distantly related birds have distinct neuronal cerebrotypes.

• Keywords
• Brain size, Cognition, Evolution, Intelligence, Number of neurons,
• MeSH
• Biological Evolution MeSH
• Species Specificity MeSH
• Galliformes * anatomy & histology   physiology   MeSH
• Brain * cytology   anatomy & histology   MeSH
• Neurons * cytology   physiology   MeSH
• Parrots * anatomy & histology   physiology   MeSH
• Cell Count MeSH
• Telencephalon cytology   MeSH
• Songbirds * anatomy & histology   physiology   MeSH
• Animals MeSH
• Check Tag
• Animals MeSH
• Publication type
• Journal Article MeSH
• Comparative Study MeSH

Mirror self-recognition (MSR) assessed by the Mark Test has been the staple test for the study of animal self-awareness. When tested in this paradigm, corvid species return discrepant results, with only the Eurasian magpies and the Indian house crow successfully passing the test so far, whereas multiple other corvid species fail. The lack of replicability of these positive results and the large divergence in applied methodologies calls into question whether the observed differences are in fact phylogenetic or methodological, and, if so, which factors facilitate the expression of MSR in some corvids. In this study, we (1) present new results on the self-recognition abilities of common ravens, (2) replicate results of azure-winged magpies, and (3) compare the mirror responses and performances in the mark test of these two corvid species with a third corvid species: carrion crows, previously tested following the same experimental procedure. Our results show interspecies differences in the approach of and the response to the mirror during the mirror exposure phase of the experiment as well as in the subsequent mark test. However, the performances of these species in the Mark Test do not provide any evidence for their ability of self-recognition. Our results add to the ongoing discussion about the convergent evolution of MSR and we advocate for consistent methodologies and procedures in comparing this ability across species to advance this discussion.

• Keywords
• Azure-winged magpies, Carrion crow, Common raven, Mirror response, Mirror-mark test, Self-awareness,
• MeSH
• Behavior, Animal physiology   MeSH
• Phylogeny MeSH
• Passeriformes * physiology   MeSH
• Crows * MeSH
• Animals MeSH
• Check Tag
• Animals MeSH
• Publication type
• Journal Article MeSH

In vertebrates, cannabinoids modulate neuroimmune interactions through two cannabinoid receptors (CNRs) conservatively expressed in the brain (CNR1, syn. CB1) and in the periphery (CNR2, syn. CB2). Our comparative genomic analysis indicates several evolutionary losses in the CNR2 gene that is involved in immune regulation. Notably, we show that the CNR2 gene pseudogenized in all parrots (Psittaciformes). This CNR2 gene loss occurred because of chromosomal rearrangements. Our positive selection analysis suggests the absence of any specific molecular adaptations in parrot CNR1 that would compensate for the CNR2 loss in the modulation of the neuroimmune interactions. Using transcriptomic data from the brains of birds with experimentally induced sterile inflammation we highlight possible functional effects of such a CNR2 gene loss. We compare the expression patterns of CNR and neuroinflammatory markers in CNR2-deficient parrots (represented by the budgerigar, Melopsittacus undulatus and five other parrot species) with CNR2-intact passerines (represented by the zebra finch, Taeniopygia guttata). Unlike in passerines, stimulation with lipopolysaccharide resulted in neuroinflammation in the parrots linked with a significant upregulation of expression in proinflammatory cytokines (including interleukin 1 beta (IL1B) and 6 (IL6)) in the brain. Our results indicate the functional importance of the CNR2 gene loss for increased sensitivity to brain inflammation.

• Keywords
• avian immunology, cannabinoid receptor pseudogenization, cannabinoid receptors, gene loss, neural inflammation, neuroimmunology,
• MeSH
• Parrots * genetics   MeSH
• Receptors, Cannabinoid MeSH
• Animals MeSH
• Check Tag
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH
• Names of Substances
• Receptors, Cannabinoid MeSH

A longstanding issue in biology is whether the intelligence of animals can be predicted by absolute or relative brain size. However, progress has been hampered by an insufficient understanding of how neuron numbers shape internal brain organization and cognitive performance. On the basis of estimations of neuron numbers for 111 bird species, we show here that the number of neurons in the pallial telencephalon is positively associated with a major expression of intelligence: innovation propensity. The number of pallial neurons, in turn, is greater in brains that are larger in both absolute and relative terms and positively covaries with longer post-hatching development periods. Thus, our analyses show that neuron numbers link cognitive performance to both absolute and relative brain size through developmental adjustments. These findings help unify neuro-anatomical measures at multiple levels, reconciling contradictory views over the biological significance of brain expansion. The results also highlight the value of a life history perspective to advance our understanding of the evolutionary bases of the connections between brain and cognition.

• MeSH
• Intelligence physiology   MeSH
• Brain physiology   MeSH
• Neurons * physiology   MeSH
• Birds * physiology   MeSH
• Organ Size MeSH
• Animals MeSH
• Check Tag
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH

SignificanceThe evolution of brain processing capacity has traditionally been inferred from data on brain size. However, similarly sized brains of distantly related species can differ in the number and distribution of neurons, their basic computational units. Therefore, a finer-grained approach is needed to reveal the evolutionary paths to increased cognitive capacity. Using a new, comprehensive dataset, we analyzed brain cellular composition across amniotes. Compared to reptiles, mammals and birds have dramatically increased neuron numbers in the telencephalon and cerebellum, which are brain parts associated with higher cognition. Astoundingly, a phylogenetic analysis suggests that as few as four major changes in neuron-brain scaling in over 300 million years of evolution pave the way to intelligence in endothermic land vertebrates.

• Keywords
• brain size, cognition, evolution, intelligence, number of neurons,
• MeSH
• Biological Evolution * MeSH
• Phylogeny MeSH
• Quantitative Trait, Heritable MeSH
• Brain cytology   physiology   MeSH
• Neurons cytology   MeSH
• Vertebrates * classification   MeSH
• Cell Count * MeSH
• Animals MeSH
• Check Tag
• Animals MeSH
• Publication type
• Journal Article MeSH

Recent studies indicate that yawning evolved as a brain cooling mechanism. Given that larger brains have greater thermolytic needs and brain temperature is determined in part by heat production from neuronal activity, it was hypothesized that animals with larger brains and more neurons would yawn longer to produce comparable cooling effects. To test this, we performed the largest study on yawning ever conducted, analyzing 1291 yawns from 101 species (55 mammals; 46 birds). Phylogenetically controlled analyses revealed robust positive correlations between yawn duration and (1) brain mass, (2) total neuron number, and (3) cortical/pallial neuron number in both mammals and birds, which cannot be attributed solely to allometric scaling rules. These relationships were similar across clades, though mammals exhibited considerably longer yawns than birds of comparable brain and body mass. These findings provide further evidence suggesting that yawning is a thermoregulatory adaptation that has been conserved across amniote evolution.

• MeSH
• Brain anatomy & histology   physiology   MeSH
• Neurons cytology   physiology   MeSH
• Birds anatomy & histology   physiology   MeSH
• Mammals anatomy & histology   physiology   MeSH
• Organ Size MeSH
• Yawning * MeSH
• Animals MeSH
• Check Tag
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH

Cerebrospinal fluid (CSF) proteins regulate neurogenesis, brain homeostasis and participate in signalling during neuroinflammation. Even though birds represent valuable models for constitutive adult neurogenesis, current proteomic studies of the avian CSF are limited to chicken embryos. Here we use liquid chromatography-tandem mass spectrometry (nLC-MS/MS) to explore the proteomic composition of CSF and plasma in adult chickens (Gallus gallus) and evolutionarily derived parrots: budgerigar (Melopsittacus undulatus) and cockatiel (Nymphicus hollandicus). Because cockatiel lacks a complete genome information, we compared the cross-species protein identifications using the reference proteomes of three model avian species: chicken, budgerigar and zebra finch (Taeniopygia guttata) and found the highest identification rates when mapping against the phylogenetically closest species, the budgerigar. In total, we identified 483, 641 and 458 unique proteins consistently represented in the CSF and plasma of all chicken, budgerigar and cockatiel conspecifics, respectively. Comparative pathways analyses of CSF and blood plasma then indicated clusters of proteins involved in neurogenesis, neural development and neural differentiation overrepresented in CSF in each species. This study provides the first insight into the proteomics of adult avian CSF and plasma and brings novel evidence supporting the adult neurogenesis in birds.

• MeSH
• Neurogenesis * MeSH
• Proteome metabolism   MeSH
• Birds * growth & development   metabolism   MeSH
• Animals MeSH
• Check Tag
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH
• Names of Substances
• Proteome MeSH

Within-species variation in the number of neurons, other brain cells and their allocation to different brain parts is poorly studied. Here, we assess these numbers in a squamate reptile, the Madagascar ground gecko (Paroedura picta). We examined adults from two captive populations and three age groups within one population. Even though reptiles exhibit extensive adult neurogenesis, intrapopulation variation in the number of neurons is similar to that in mice. However, the two populations differed significantly in most measures, highlighting the fact that using only one population can underestimate within-species variation. There is a substantial increase in the number of neurons and decrease in neuronal density in adult geckos relative to hatchlings and an increase in the number of neurons in the telencephalon in fully grown adults relative to sexually mature young adults. This finding implies that adult neurogenesis does not only replace worn out but also adds new telencephalic neurons in reptiles during adulthood. This markedly contrasts with the situation in mammals, where the number of cortical neurons declines with age.

• Keywords
• brain size, cellular scaling rules, evolution, intraspecific variation, number of glial cells, number of neurons,
• MeSH
• Lizards * MeSH
• Brain MeSH
• Mice MeSH
• Neurons MeSH
• Telencephalon MeSH
• Animals MeSH
• Check Tag
• Mice MeSH
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH
• Geographicals
• Madagascar MeSH

The stereotyped features of brain structure, such as the distribution, morphology and connectivity of neuronal cell types across brain areas, are those most likely to explain the remarkable capacity of the brain to process information and govern behaviors. Recent advances in anatomical methods, including the simple but versatile isotropic fractionator and several whole-brain labeling, clearing and microscopy methods, have opened the door to an exciting new era in comparative brain anatomy, one that has the potential to transform our understanding of the brain structure-function relationship by representing the evolution of brain complexity in quantitative anatomical features shared across species and species-specific or clade-specific. Here we discuss these methods and their application to mapping brain cell count and cell type distributions-two particularly powerful neural correlates of vertebrate cognitive and behavioral capabilities.

• MeSH
• Species Specificity MeSH
• Brain Mapping * MeSH
• Brain * MeSH
• Neurons MeSH
• Cell Count MeSH
• Animals MeSH
• Check Tag
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH
• Review MeSH
• Research Support, N.I.H., Extramural MeSH

Neurons are the basic computational units of the brain, but brain size is the predominant surrogate measure of brain functional capacity in comparative and cognitive neuroscience. This approach is based on the assumption that larger brains harbor higher numbers of neurons and their connections, and therefore have a higher information-processing capacity. However, recent studies have shown that brain mass may be less strongly correlated with neuron counts than previously thought. Till now, no experimental test has been conducted to examine the relationship between evolutionary changes in brain size and the number of brain neurons. Here, we provide such a test by comparing neuron number in artificial selection lines of female guppies (Poecilia reticulata) with >15% difference in relative brain mass and numerous previously demonstrated cognitive differences. Using the isotropic fractionator, we demonstrate that large-brained females have a higher overall number of neurons than small-brained females, but similar neuronal densities. Importantly, this difference holds also for the telencephalon, a key region for cognition. Our study provides the first direct experimental evidence that selection for brain mass leads to matching changes in number of neurons and shows that brain size evolution is intimately linked to the evolution of neuron number and cognition.

• Keywords
• Artificial selection, brain size, cognition, isotropic fractionator, number of neurons,
• MeSH
• Biological Evolution MeSH
• Cognition MeSH
• Models, Neurological MeSH
• Brain physiology   MeSH
• Neurons physiology   MeSH
• Selection, Genetic * MeSH
• Organ Size MeSH
• Poecilia genetics   physiology   MeSH
• Animals MeSH
• Check Tag
• Female MeSH
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH