Telencephalon Size Research Articles

Eye size does not change with artificial selection on relative telencephalon size in guppies (Poecilia reticulata).

Variation in eye size is sometimes closely associated with brain morphology. Visual information, detected by the retina, is transferred to the optic tectum to coordinate eye and body movements towards stimuli, and thereafter distributed into other brain regions for further processing. The telencephalon is an important visual processing region in many vertebrate species and a highly developed region in visually dependent species. Yet, the existence of a coevolutionary relationship between telencephalon size and eye size remains relatively unknown. Here, we use male and female guppies artificially selected for small- and large-relative-telencephalon-size to test if artificial selection on telencephalon size results in changes in eye size. In addition, we performed an optomotor test as a proxy for visual acuity. We found no evidence that eye size changes with artificial selection on telencephalon size. Eye size was similar in both absolute and relative terms between the two selection regimes, but was larger in females. This is most likely because of the larger body size in females, but it could also reflect their greater need for visual capacity due to sex-specific differences in foraging and mating behaviour. Although the optomotor response was stronger in guppies with a larger telencephalon, we found no evidence for differences in visual acuity between the selection regimes. Our study suggests that eye size and visual perception in guppies does not change rapidly with strong artificial selection on telencephalon size.

Relative telencephalon size does not affect collective motion in the guppy (Poecilia reticulata).

Collective motion is common across all animal taxa, from swarming insects to schools of fish. The collective motion requires intricate behavioral integration among individuals, yet little is known about how evolutionary changes in brain morphology influence the ability for individuals to coordinate behavior in groups. In this study, we utilized guppies that were selectively bred for relative telencephalon size, an aspect of brain morphology that is normally associated with advanced cognitive functions, to examine its role in collective motion using an open-field assay. We analyzed high-resolution tracking data of same-sex shoals consisting of 8 individuals to assess different aspects of collective motion, such as alignment, attraction to nearby shoal members, and swimming speed. Our findings indicate that variation in collective motion in guppy shoals might not be strongly affected by variation in relative telencephalon size. Our study suggests that group dynamics in collectively moving animals are likely not driven by advanced cognitive functions but rather by fundamental cognitive processes stemming from relatively simple rules among neighboring individuals.

A Comparison of Telencephalon Composition among Chickens, Junglefowl, and Wild Galliforms

Introduction: Domestication is the process of modifying animals for human benefit through selective breeding in captivity. One of the traits that often diverges is the size of the brain and its constituent regions; almost all domesticated species have relatively smaller brains and brain regions than their wild ancestors. Although the effects of domestication on the brain have been investigated across a range of both mammal and bird species, almost nothing is known about the neuroanatomical effects of domestication on the world’s most common bird: the chicken (Gallus gallus). Methods: We compared the quantitative neuroanatomy of the telencephalon of white leghorn chickens with red junglefowl, their wild counterpart, and several wild galliform species. We focused specifically on the telencephalon because telencephalic regions typically exhibit the biggest differences in size in domesticate-wild comparisons. Results: Relative telencephalon size was larger in chickens than in junglefowl and ruffed grouse (Bonasa umbellus). The relative size of telencephalic regions did not differ between chickens and junglefowl, but did differ in comparison with ruffed grouse. Ruffed grouse had larger hyperpallia and smaller entopallial, nidopallial, and striatal volumes than chickens and junglefowl. Multivariate analyses that included an additional three wild grouse species corroborated these findings: chicken and junglefowl have relatively larger nidopallial and striatal volumes than grouse. Conversely, the mesopallial and hyperpallial volumes tended to be relatively smaller in chickens and junglefowl. Conclusion: From this suite of comparisons, we conclude that chickens do not follow a pattern of widespread decreases in telencephalic region sizes that is often viewed as typical of domestication. Instead, chickens have undergone a mosaic of changes with some regions increasing and others decreasing in size, and there are few differences between chickens and junglefowl.

No Sex-Specific Effects of Artificial Selection for Relative Telencephalon Size during Detour Learning and Spatial Discrimination in Guppies (Poecilia reticulata)

Over recent decades, substantial research has focused on fish cognitive evolution to increase our understanding of the evolution of the enormous diversity of cognitive abilities that exists in fishes. One important but understudied aspect of cognitive evolution is sexual dimorphism in cognitive abilities. Sex-specific variation in brain region morphology has been proposed to be an important mechanism in this context. However, it is also common to find sex-specific variation in behavior and cognition without associated differences in brain morphology among the sexes. The telencephalon is the major cognitive center in the vertebrate brain and variation in telencephalon size has been associated with variation in cognition. Here, we utilize recently developed guppy artificial selection lines with ca. 10% differences in relative telencephalon size to investigate whether similar responses to selection of the size of this region may affect cognitive abilities differently in males and females. To that end, we compared two ecologically relevant aspects of cognition, detour learning and binary spatial discrimination. We tested the significance of the interaction between telencephalon size and sex, and we found no sex-specific effects of evolutionary increases in telencephalon size in the cognitive abilities tested. This study indicates that no clear cognitive sex-specific effects occur in response to rapid selection of telencephalon size. We suggest that future research on sexual dimorphism in cognitive abilities in fish could use various cognitive tests and examine telencephalic sub-regions to gain a more comprehensive understanding of their evolution.

Experimental expansion of relative telencephalon size improves the main executive function abilities in guppy.

Executive functions are a set of cognitive control processes required for optimizing goal-directed behavior. Despite more than two centuries of research on executive functions, mostly in humans and nonhuman primates, there is still a knowledge gap in what constitutes the mechanistic basis of evolutionary variation in executive function abilities. Here, we show experimentally that size changes in a forebrain structure (i.e. telencephalon) underlie individual variation in executive function capacities in a fish. For this, we used male guppies (Poecilia reticulata) issued from artificial selection lines with substantial differences in telencephalon size relative to the rest of the brain. We tested fish from the up- and down-selected lines not only in three tasks for the main core executive functions: cognitive flexibility, inhibitory control, and working memory, but also in a basic conditioning test that does not require executive functions. Individuals with relatively larger telencephalons outperformed individuals with smaller telencephalons in all three executive function assays but not in the conditioning assay. Based on our findings, we propose that the telencephalon is the executive brain in teleost fish. Together, it suggests that selective enlargement of key brain structures with distinct functions, like the fish telencephalon, is a potent evolutionary pathway toward evolutionary enhancement of advanced cognitive abilities in vertebrates.

Brain morphology correlates of learning and cognitive flexibility in a fish species (Poecilia reticulata).

Determining how variation in brain morphology affects cognitive abilities is important to understand inter-individual variation in cognition and, ultimately, cognitive evolution. Yet, despite many decades of research in this area, there is surprisingly little experimental data available from assays that quantify cognitive abilities and brain morphology in the same individuals. Here, we tested female guppies (Poecilia reticulata) in two tasks, colour discrimination and reversal learning, to evaluate their learning abilities and cognitive flexibility. We then estimated the size of five brain regions (telencephalon, optic tectum, hypothalamus, cerebellum and dorsal medulla), in addition to relative brain size. We found that optic tectum relative size, in relation to the rest of the brain, correlated positively with discrimination learning performance, while relative telencephalon size correlated positively with reversal learning performance. The other brain measures were not associated with performance in either task. By evaluating how fast learning occurs and how fast an animal adjusts its learning rules to changing conditions, we find support for that different brain regions have distinct functional correlations at the individual level. Importantly, telencephalon size emerges as an important neural correlate of higher executive functions such as cognitive flexibility. This is rare evidence supporting the theory that more neural tissue in key brain regions confers cognitive benefits.

Latent effect of larval rearing environment on post-metamorphic brain growth in an anuran amphibian

Early development is highly susceptible to environmental influence. We evaluated the role of larval visual environment on brain morphology plasticity in late larval and juvenile stages of Bombina orientalis, an anuran amphibian changing from an aquatic to a terrestrial habitat after metamorphosis. Manipulation of the visual environment was achieved by rearing larvae in normal and darkened water. The juveniles were exposed to normal lighting conditions after metamorphosis, allowing to assess if plastic effects persisted or emerged after metamorphosis. The darkness treatment accelerated development before slowing it down substantially, allowing controls to metamorphose earlier. Although larvae reared in darkened water had the same relative brain size as controls by the end of the larval period, juveniles that had been reared in darkened water as larvae had brains that were 14.4% smaller than juveniles that had been reared under control conditions. Conversely, relative telencephalon size was 6.7% larger in juveniles previously reared in darkened water compared with controls, again with no effect of darkened water seen by the end of the larval period. Unlike the latent effects seen on whole brain and telencephalon size, relative size of the optic tectum was significantly smaller in both larvae and juveniles exposed to the darkened water treatment. Therefore, the effects of visual restriction on juvenile brain form were a combination of latent (whole brain and telencephalon) and carry-over (optic tectum) developmental effects.

Artificial mosaic brain evolution of relative telencephalon size improves inhibitory control abilities in the guppy (Poecilia reticulata).

Mosaic brain evolution, the change in the size of separate brain regions in response to selection on cognitive performance, is an important idea in the field of cognitive evolution. However, untill now, most of the data on how separate brain regions respond to selection and their cognitive consequences stem from comparative studies. To experimentally investigate the influence of mosaic brain evolution on cognitive ability, we used male guppies artificially selected for large and small telencephalons relative to the rest of the brain. Here, we tested an important aspect of executive cognitive ability using a detour task. We found that males with larger telencephalons outperformed males with smaller telencephalons. Fish with larger telencephalons showed faster improvement in performance during detour training and were more successful in reaching the food reward without touching the transparent barrier (i.e., through correct detouring) during the test phase. Together, our findings provide the first experimental evidence showing that evolutionary enlargement of relative telencephalon size confers cognitive benefits, supporting an important role for mosaic brain evolution during cognitive evolution.

Rapid mosaic brain evolution under artificial selection for relative telencephalon size in the guppy (Poecilia reticulata).

The mosaic brain evolution hypothesis, stating that brain regions can evolve relatively independently during cognitive evolution, is an important idea to understand how brains evolve with potential implications even for human brain evolution. Here, we provide the first experimental evidence for this hypothesis through an artificial selection experiment in the guppy (Poecilia reticulata). After four generations of selection on relative telencephalon volume (relative to brain size), we found substantial changes in telencephalon size but no changes in other regions. Further comparisons revealed that up-selected lines had larger telencephalon, while down-selected lines had smaller telencephalon than wild Trinidadian populations. Our results support that independent evolutionary changes in specific brain regions through mosaic brain evolution can be important facilitators of cognitive evolution.

Does brain size affect mate choice? An experimental examination in pygmy halfbeaks.

Choosing a mate is one of the most important decisions in an animal’s lifetime. Female mate choice is often guided by the presence or intensity of male sexual ornaments, which must be integrated and compared among potential mates. Individuals with greater cognitive abilities may be better at evaluating and comparing sexual ornaments, even when the difference in ornaments is small. While brain size is often used as a proxy for cognitive ability, its effect on mate choice has rarely been investigated. Here, we investigate the effect of brain size on mate preferences in the pygmy halfbeak Dermogenys collettei, a small freshwater fish that forms mixed-sex shoals where mating takes place. Pygmy halfbeaks are ideal models as their semi-transparent heads allow for external brain measurements. After validating the use of external measurements as a proxy for internal brain size, we presented females with large or small brains (relative to body length) with two males that had either a large or small difference in sexual ornamentation (measured by the total area of red coloration). Unexpectedly, neither total relative brain size nor relative telencephalon size affected any measured aspect of mate preference. However, the difference in male sexual ornamentation did affect preference, with females preferring males with a smaller area of red coloration when the difference in ornaments was large. This study highlights the complexities of mate choice and the importance of considering a range of stimuli when examining mate preferences.

Rab11fip5 regulates telencephalon development via ephrinB1 recycling.

ABSTRACTRab11 family-interacting protein 5 (Rab11fip5) is an adaptor protein that binds to the small GTPase Rab11, which has an important function in endosome recycling and trafficking of cellular proteins to the plasma membrane. Rab11fip5 is involved in many cellular processes, such as cytoskeleton rearrangement, iron uptake and exocytosis in neuroendocrine cells, and is also known as a candidate gene for autism-spectrum disorder. However, the role of Rab11fip5 during early embryonic development is not clearly understood. In this study, we identified Rab11fip5 as a protein that interacts with ephrinB1, a transmembrane ligand for Eph receptors. The PDZ binding motif in ephrinB1 and the Rab-binding domain in Rab11fip5 are necessary for their interaction in a complex. EphrinB1 and Rab11fip5 display overlapping expression in the telencephalon of developing amphibian embryos. The loss of Rab11fip5 function causes a reduction in telencephalon size and a decrease in the expression level of ephrinB1. Moreover, morpholino oligonucleotide-mediated knockdown of Rab11fip5 decreases cell proliferation in the telencephalon. The overexpression of ephrinB1 rescues these defects, suggesting that ephrinB1 recycling by the Rab11/Rab11fip5 complex is crucial for proper telencephalon development.

Predation pressure shapes brain anatomy in the wild

There is remarkable diversity in brain anatomy among vertebrates and evidence is accumulating that predatory interactions are crucially important for this diversity. To test this hypothesis, we collected female guppies (Poecilia reticulata) from 16 wild populations and related their brain anatomy to several aspects of predation pressure in this ecosystem, such as the biomass of the four major predators of guppies (one prawn and three fish species), and predator diversity (number of predatory fish species in each site). We found that populations from localities with higher prawn biomass had relatively larger telencephalon size as well as larger brains. Optic tectum size was positively associated with one of the fish predator’s biomass and with overall predator diversity. However, both olfactory bulb and hypothalamus size were negatively associated with the biomass of another of the fish predators. Hence, while fish predator occurrence is associated with variation in brain anatomy, prawn occurrence is associated with variation in brain size. Our results suggest that cognitive challenges posed by local differences in predator communities may lead to changes in prey brain anatomy in the wild.

Brain morphology of the threespine stickleback (Gasterosteus aculeatus) varies inconsistently with respect to habitat complexity: A test of the Clever Foraging Hypothesis.

The Clever Foraging Hypothesis asserts that organisms living in a more spatially complex environment will have a greater neurological capacity for cognitive processes related to spatial memory, navigation, and foraging. Because the telencephalon is often associated with spatial memory and navigation tasks, this hypothesis predicts a positive association between telencephalon size and environmental complexity. The association between habitat complexity and brain size has been supported by comparative studies across multiple species but has not been widely studied at the within‐species level. We tested for covariation between environmental complexity and neuroanatomy of threespine stickleback (Gasterosteus aculeatus) collected from 15 pairs of lakes and their parapatric streams on Vancouver Island. In most pairs, neuroanatomy differed between the adjoining lake and stream populations. However, the magnitude and direction of this difference were inconsistent between watersheds and did not covary strongly with measures of within‐site environmental heterogeneity. Overall, we find weak support for the Clever Foraging Hypothesis in our study.

Brain size evolution in the frog Fejervarya limnocharis supports neither the cognitive buffer nor the expensive brain hypothesis

AbstractVariable environmental conditions affect brain size evolution. To explain inter‐ and intraspecific brain size variation among vertebrates, two adaptive hypotheses – the expensive brain hypothesis (EBH) and the cognitive buffer hypothesis (CBH) – have been proposed. The EBF proposes that relative brain size is reduced in animals that experience longer periods of low food availability in fluctuating environments. Alternatively, the CBH states that a major advantage of a relatively large brain is to enhance cognitive abilities in fluctuating environments. We found considerable variation in brain size and the sizes of brain structures in the Asian grass frog (Fejervarya limnocharis) among populations. Inconsistent with the predictions of the EBH and the CBH, individuals living in habitats with medium variation in temperatures and length of growth seasons at middle latitude and/or altitude had a relatively large brain, suggesting that environmental habitat‐induced energetic constraints do not play an important role in shaping brain size evolution. Moreover, we also detected a significant intersexual difference in telencephalon size. Specifically, none of relative size of brain structures was correlated with variation in temperature and growth season related to environmental habitats, although latitude strongly affected the relative sizes of telencephalon and cerebellum. Our findings decline both the EBH and the CBH to explain brain size variation in the Asian grass frog. Furthermore, our results also suggest the sex‐specificity of the environmental‐induced brain size evolution in adult frogs associated with reproductive behaviors and spawning site selection.

Phylogenetic ANCOVA: Estimating Changes in Evolutionary Rates as Well as Relationships between Traits.

We present a new phylogenetic comparative method-phylogenetic analysis of covariance (PANCOVA)-that uses interspecific data and a phylogeny to estimate the effects of major events on both the rate of phenotypic evolution and the association between traits. It could be used, for example, to model the impact of a key innovation, colonization of a new habitat, or environmental change. The approach is optimized with maximum likelihood and is formulated under the familiar phylogenetic generalized least squares framework, which is flexible and easily extended to incorporate other factors and parameters. As an example, we explore the relationship between parental investment and relative telencephalon size in birds and contrast the results of PANCOVA with those from other phylogenetic comparative methods.

Evolution of anuran brains: disentangling ecological and phylogenetic sources of variation.

Variation in ecological selection pressures has been implicated to explain variation in brain size and architecture in fishes, birds and mammals, but little is known in this respect about amphibians. Likewise, the relative importance of constraint vs. mosaic hypotheses of brain evolution in explaining variation in brain size and architecture remains contentious. Using phylogenetic comparative methods, we studied interspecific variation in brain size and size of different brain parts among 43 Chinese anuran frogs and explored how much of this variation was explainable by variation in ecological factors (viz. habitat type, diet and predation risk). We also evaluated which of the two above-mentioned hypotheses best explains the observed patterns. Although variation in brain size explained on average 80.5% of the variation in size of different brain parts (supporting the constraint hypothesis), none of the three ecological factors were found to explain variation in overall brain size. However, habitat and diet type explained a significant amount of variation in telencephalon size, as well in three composite measures of brain architecture. Likewise, predation risk explained a significant amount of variation in bulbus olfactorius and optic tecta size. Our results show that evolution of anuran brain accommodates features compatible with both constraint (viz. strong allometry among brain parts) and mosaic (viz. independent size changes in response to ecological factors in certain brain parts) models of brain size evolution.

The Mammalian DM Domain Transcription Factor Dmrta2 Is Required for Early Embryonic Development of the Cerebral Cortex

Development of the mammalian telencephalon is precisely organized by a combination of extracellular signaling events derived from signaling centers and transcription factor networks. Using gene expression profiling of the developing mouse dorsal telencephalon, we found that the DM domain transcription factor Dmrta2 (doublesex and mab-3-related transcription factor a2) is involved in the development of the dorsal telencephalon. Consistent with its medial-high/lateral-low expression pattern in the dorsal telencephalon, Dmrta2 null mutants demonstrated a dramatic reduction in medial cortical structures such as the cortical hem and the choroid plexus, and a complete loss of the hippocampus. In this mutant, the dorsal telencephalon also showed a remarkable size reduction, in addition to abnormal cell cycle kinetics and defective patterning. In contrast, a conditional Dmrta2 deletion in the telencephalon, which was accomplished after entry into the neurogenic phase, resulted in only a slight reduction in telencephalon size and normal patterning. We also found that Dmrta2 expression was decreased by a dominant-negative Tcf and was increased by a stabilized β-catenin form. These data suggest that Dmrta2 plays pivotal roles in the early development of the telencephalon via the formation of the cortical hem, a source of Wnts, and also in the maintenance of neural progenitors as a downstream of the Wnt pathway.

Phenotypic plasticity of the threespine stickleback Gasterosteus aculeatus telencephalon in response to experience in captivity

Abstract Threespine stickleback were used to examine phenotypic plasticity of telencephala in relation to inferred ecology. Fish from derived, allopatric, freshwater populations were sampled from three shallow, structurally complex lakes with ben-thic-foraging stickleback (benthics) and from three deep, structurally simple lakes with planktivores (limnetics). The telencepha-lon of specimens preserved immediately after capture (field-preserved), field-caught fish held in aquaria for 90 days (lab-held), and lab-bred fish from crosses and raised in aquaria were compared. Field-preserved sea-run (ancestral) stickleback were collected from two separate sites, and parents of lab-bred sea-run stickleback were collected from one of these sites. In field-preserved and lab-held fish, the telencephala of limnetics exhibited triangular dorsal shape, while those of benthics and sea-run fish had rounder shapes. No such pattern was detected in lab-bred fish. Within each treatment type, benthics had larger relative telencephalon sizes, using overall brain size as the covariate, than limnetics. Among field-preserved samples, sea-run fish had smaller telencephalon sizes than lake fish. Intra-population analyses of lake samples showed that field-preserved fish consistently had larger relative telencephalon sizes than lab-bred fish. The opposite was true of the sea-run population. In a separate study using one benthic population and one limnetic population, samples were preserved in the field immediately or held in the lab for 30, 60, and 90 days before they were sacrificed. In both populations, the telencephalon shapes of lab-held fish were similar to those of field-preserved fish but became progressively more like lab-bred ones over 90 days. In contrast, relative telencephalon size decreased dramatically by 30 days after which there was little change. In freshwater threespine stickleback, the telencephalon exhibits considerable phenotypic plasticity, which was probably present in the ancestor.

Variation of telencephalon morphology of the threespine stickleback (Gasterosteus aculeatus) in relation to inferred ecology

We tested the hypothesis that increased telencephalon size has evolved in threespine stickleback fish (Gasterosteus aculeatus) from structurally complex habitats using field-caught samples from one sea-run (ancestral) and 18 ecologically diverse freshwater (descendant) populations. Freshwater habitats ranged from shallow, structurally complex lakes with benthic-foraging stickleback (benthics), to deeper, structurally simple lakes in which stickleback depend more heavily on plankton for prey (generalists). Contrary to our expectations, benthics had smaller telencephala than generalists, but the shape of the telencephalon of the sea-run and benthic populations were more convex laterally. Convex telencephalon shape may indicate enlargement of the dorsolateral region, which is homologous with the tetrapod hippocampus. Telencephalon morphology is also sexually dimorphic, with larger, less convex telencephala in males. Freshwater stickleback from structurally complex habitats have retained the ancestral telencephalon morphology, but populations that feed more in open habitats on plankton have evolved larger, laterally concave telencephala.

Comparative Analyses of Song Complexity and Song-Control Nuclei in Fourteen Oscine Species

Most studies on the relationship between measures of song behavior and the sizes of song control nuclei have focused on one or two oscine species, and often show inconsistent results. To address this issue, we first measured four variables for song complexity, i.e., song repertoire size, syllable repertoire size, the mean number of syllables per phrase (MNS) and the number of syllables in the longest phrase (NSLP), and the sizes of three song control nuclei, i.e., HVC, RA (the robust nucleus of the arcopallium), and Area X in 14 oscine species from eight families. To tackle the problem of statistical non-independence that probably existed among the closely related species, we reconstructed the phylogeny of the species studied using mitochondrial cytochrome b DNA sequences from GenBank. By using the methods adopted in most previous reports, we tested the relationship between song complexity and the sizes of the song control nuclei. We found that: 1) the absolute sizes of RA and Area X, but not of HVC, were positively correlated to the three measures of song complexity, but that only the residual size of RA in regard to telencephalon size was significantly correlated to the song measures; 2) independent contrasts analysis showed RA and Area X to be significantly associated with NSLP. Our results indicated that the relationship between song behavior and its neural structures varied among song nuclei, suggesting that each song control nucleus may play a different role in song behavior.