Input and Output Connections of the Crow Nidopallium Caudolaterale.

The avian telencephalic structure nidopallium caudolaterale (NCL) functions as an analog to the mammalian prefrontal cortex. In crows, corvid songbirds, it plays a crucial role in higher cognitive and executive functions. These functions rely on the NCL's extensive telencephalic connections. However, systematic investigations into the brain-wide connectivity of the NCL in crows or other songbirds are lacking. Here, we studied its input and output connections by injecting retrograde and anterograde tracers into the carrion crow NCL. Our results, mapped onto a published carrion crow brain atlas, confirm NCL multisensory connections and extend prior pigeon findings by identifying a novel input from the hippocampal formation. Furthermore, we analyze crow NCL efferent projections to the arcopallium and report newly identified arcopallial neurons projecting bilaterally to the NCL. These findings help to clarify the role of the NCL as central executive hub in the corvid songbird brain.

Molecular biology of serotonergic systems in avian brains.

Serotonin (5-hydroxytryptamine, 5-HT) is a phylogenetically conserved neurotransmitter and modulator. Neurons utilizing serotonin have been identified in the central nervous systems of all vertebrates. In the central serotonergic system of vertebrate species examined so far, serotonergic neurons have been confirmed to exist in clusters in the brainstem. Although many serotonin-regulated cognitive, behavioral, and emotional functions have been elucidated in mammals, equivalents remain poorly understood in non-mammalian vertebrates. The purpose of this review is to summarize current knowledge of the anatomical organization and molecular features of the avian central serotonergic system. In addition, selected key functions of serotonin are briefly reviewed. Gene association studies between serotonergic system related genes and behaviors in birds have elucidated that the serotonergic system is involved in the regulation of behavior in birds similar to that observed in mammals. The widespread distribution of serotonergic modulation in the central nervous system and the evolutionary conservation of the serotonergic system provide a strong foundation for understanding and comparing the evolutionary continuity of neural circuits controlling corresponding brain functions within vertebrates. The main focus of this review is the chicken brain, with this type of poultry used as a model bird. The chicken is widely used not only as a model for answering questions in developmental biology and as a model for agriculturally useful breeding, but also in research relating to cognitive, behavioral, and emotional processes. In addition to a wealth of prior research on the projection relationships of avian brain regions, detailed subdivision similarities between avian and mammalian brains have recently been identified. Therefore, identifying the neural circuits modulated by the serotonergic system in avian brains may provide an interesting opportunity for detailed comparative studies of the function of serotonergic systems in mammals.

How smart dinosaurs?

This commentary discusses the main points made in Reiner's article on the prospect that some theropod dinosaurs could have given rise to a lineage that achieved a human level of intelligence, and those made in Herculano-Houzel's article on the potentially monkey-like numbers of neurons in the pallium of large theropods, and the implications of this for their intelligence.

Could theropod dinosaurs have evolved to a human level of intelligence?

Noting that some theropod dinosaurs had large brains, large grasping hands, and likely binocular vision, paleontologist Dale Russell suggested that a branch of these dinosaurs might have evolved to a human intelligence level, had dinosaurs not become extinct. I offer reasons why the likely pallial organization in dinosaurs would have made this improbable, based on four assumptions. First, it is assumed that achieving human intelligence requires evolving an equivalent of the about 200 functionally specialized cortical areas characteristic of humans. Second, it is assumed that dinosaurs had an avian nuclear type of pallial organization, in contrast to the mammalian cortical organization. Third, it is assumed that the interactions between the different neuron types making up an information processing unit within pallium are critical to its role in analyzing information. Finally, it is assumed that increasing axonal length between the neuron sets carrying out this operation impairs its efficacy. Based on these assumptions, I present two main reasons why dinosaur pallium might have been unable to add the equivalent of 200 efficiently functioning cortical areas. First, a nuclear pattern of pallial organization would require increasing distances between the neuron groups corresponding to the separate layers of any given mammalian cortical area, as more sets of nuclei equivalent to a cortical area are interposed between the existing sets, increasing axon length and thereby impairing processing efficiency. Second, because of its nuclear organization, dinosaur pallium could not reduce axon length by folding to bring adjacent areas closer together, as occurs in cerebral cortex.

The brain-specific upregulation of CARD11 in response to avian brain-neurotropic virus infection serves as a potential biomarker.

Avian neurotropic viruses are critical problems in poultry industry causing severe central nervous system (CNS) damage with neuroinvasive and neurovirulence properties. Biomarker of neurotropic viral intracranial invasion is of great application value for the diagnosis, but that of avian neurotropic viruses remains elusive. Previously, we found that chicken caspase recruitment domain family, member 11 (CARD11) was only upregulated in virulent Newcastle disease virus-infected chickens and in chicken primary neuronal cells. In this study, CARD11 was systemically expressed in chickens and pigeons detected by absolute qPCR and immunohistochemical (IHC) assay. After virus challenging, only avian neurotropic viruses (avian encephalomyelitis virus [AEV] and pigeon paramyxovirus type 1 [PPMV-1]) except Marek's disease virus (MDV) can invade brain and cause pathological changes. The relative mRNA expression of CARD11 was brain-upregulated in AEV- or PPMV-1-infected animals, rather than MDV and non-neurotropic viruses (fowl adenovirus serotype 4 [FAdV-4] and infectious bronchitis virus [IBV]). Similarly, the protein expression of CARD11 was only upregulated in the cerebra and cerebella infected by avian brain-neurotropic virus using IHC assay. And there were no correlations between the change level of CARD11 and viral load. Our preliminary data suggested that avian CARD11 may be a potential brain biomarker for avian brain-neurotropic virus invasion.

Regional Patterning of Adult Neurogenesis in the Homing Pigeon's Brain.

In the avian brain, adult neurogenesis has been reported in the telencephalon of several species, but the functional significance of this trait is still ambiguous. Homing pigeons (Columba livia f.d.) are well-known for their navigational skills. Their brains are functionally adapted to homing with, e.g., larger hippocampi. So far, no comprehensive mapping of adult neuro- and gliogenesis or studies of different developmental neuronal stages in the telencephalon of homing pigeons exists, although comprehensive analyses in various species surely will result in a higher understanding of the functional significance of adult neurogenesis. Here, adult, free flying homing pigeons were treated with 5-bromo-deoxyuridine (BrdU) to label adult newborn cells. Brains were dissected and immunohistochemically processed with several markers (GFAP, Sox2, S100ß, Tbr2, DCX, Prox1, Ki67, NeuN, Calbindin, Calretinin) to study different stages of adult neurogenesis in a quantitative and qualitative way. Therefore, immature and adult newborn neurons and glial cells were analyzed along the anterior-posterior axis. The analysis proved the existence of different neuronal maturation stages and showed that immature cells, migrating neurons and adult newborn neurons and glia were widely and regionally unequally distributed. Double- and triple-labelling with developmental markers allowed a stage classification of adult neurogenesis in the pigeon brain (1: continuity of stem cells/proliferation, 2: fate specification, 3: differentiation/maturation, 4: integration). The most adult newborn neurons and glia were found in the intercalated hyperpallium (HI) and the hippocampal formation (HF). The highest numbers of immature (DCX+) cells were detected in the nidopallium (N). Generally, the number of newborn glial cells exceeded the number of newborn neurons. Individual structures (e.g., HI, N, and HF) showed further variations along the anterior-posterior axis. Our qualitative classification and the distribution of maturing cells in the forebrain support the idea that there is a functional specialization, respectively, that there is a link between brain-structure and function, species-specific requirements and adult neurogenesis. The high number of immature neurons also suggests a high level of plasticity, which points to the ability for rapid adaption to environmental changes through additive mechanisms. Furthermore, we discuss a possible influence of adult neurogenesis on spatial cognition.

Possible link between brain size and flight mode in birds: Does soaring ease the energetic limitation of the brain?

Elucidating determinants of interspecies variation in brain size has been a long-standing challenge in cognitive and evolutionary ecology. As the brain is an energetically expensive organ, energetic tradeoffs among organs are considered to play a key role in brain size evolution. This study examined the tradeoff between the brain and locomotion in birds by testing the relationship between brain size, flight modes with different energetic costs (flapping and soaring), and migratory behavior, using published data on the whole-brain mass of 2242 species. According to comparative analyses considering phylogeny and body mass, soarers, who can gain kinetic energy from wind shear or thermals and consequently save flight costs, have larger brains than flappers among migratory birds. Meanwhile, the brain size difference was not consistent in residents, and the size variation appeared much larger than that in migrants. In addition, the brain size of migratory birds was smaller than that of resident birds among flappers, whereas this property was not significant in soarers. Although further research is needed to draw a definitive conclusion, these findings provide further support for the energetic tradeoff of the brain with flight and migratory movements in birds and advance the idea that a locomotion mode with lower energetic cost could be a driver of encephalization during the evolution of the brain.

Telencephalic regulation of the HPA axis in birds.

The hypothalamo-pituitary-adrenal (HPA) axis is one of the major output systems of the vertebrate stress response. It controls the release of cortisol or corticosterone from the adrenal gland. These hormones regulate a range of processes throughout the brain and body, with the main function of mobilizing energy reserves to improve coping with a stressful situation. This axis is regulated in response to both physical (e.g., osmotic) and psychological (e.g., social) stressors. In mammals, the telencephalon plays an important role in the regulation of the HPA axis response in particular to psychological stressors, with the amygdala and part of prefrontal cortex stimulating the stress response, and the hippocampus and another part of prefrontal cortex inhibiting the response to return it to baseline. Birds also mount HPA axis responses to psychological stressors, but much less is known about the telencephalic areas that control this response. This review summarizes which telencephalic areas in birds are connected to the HPA axis and are known to respond to stressful situations. The conclusion is that the telencephalic control of the HPA axis is probably an ancient system that dates from before the split between sauropsid and synapsid reptiles, but more research is needed into the functional relationships between the brain areas reviewed in birds if we want to understand the level of this conservation.

Central prolactin receptor distribution and pSTAT5 activation patterns in breeding and non-breeding zebra finches (Taeniopygia guttata).

The hormone prolactin has many diverse functions across taxa such as osmoregulation, metabolism, and reproductive behavior. In ring doves, central prolactin action is important for parental care and feeding behavior. However, there is a considerable lack of information on the distribution of the prolactin receptor (PRLR) in the avian CNS to test the hypothesis that prolactin mediates these and other functions in other birds. In order to advance this research, we collected brains from breeding and non-breeding zebra finches to map the PRLR distribution using immunohistochemistry. We found PRLRs are distributed widely across the brain, both in hypothalamic sites known to regulate parental care and feeding, but also in many non-hypothalamic sites, including the tectofugal visual pathway, song system regions, reward associated areas, and pallium. This raises the possibility that prolactin has other functions throughout the brain that are not necessarily related to feeding or parental care. In addition, we also stained brains for pSTAT5, a transcription factor which is expressed when the PRLR is activated and is used as a marker for PRLR activity. We found several notable differences in pSTAT5 activity due to the breeding state of the animal, in both directions, further supporting the hypothesis that prolactin has many diverse functions in the brain both within and outside times of breeding. Together, this study represents the first essential step to inform the design of causative studies which manipulate PRLR-expressing cells to test their role in a wide variety of behaviors and other physiological functions.

Cell Proliferation in the Adult Chicken Hippocampus Correlates With Individual Differences in Time Spent in Outdoor Areas and Tonic Immobility.

Access to outdoor areas is provided as a means of enhancing welfare in commercial systems for laying hens (Gallus gallus domesticus), but substantial individual differences exist in their proportional use. Baseline cell proliferation levels of Adult Hippocampal Neurogenesis (AHN) have been associated with individual differences in reactive vs. proactive coping style, and in both mammals and birds, AHN is upregulated by positive experiences including environmental enrichment and exercise. We thus sought to explore whether individual differences in use of outdoor areas and in tonic immobility responses (indicative of fearfulness) were associated with hippocampal cell proliferation and neuronal differentiation. Radio frequency identification technology was used to track the ranging behavior of 440 individual focal hens within a commercially-relevant system over a 72-days period, after which tonic immobility durations were measured. Following hippocampal tissue collection from 58 focal hens, proliferation and neuronal differentiation were measured through quantitative PCR for proliferating cell nuclear antigen (PCNA) and doublecortin mRNA, respectively. Individual differences in tonic immobility duration positively correlated with PCNA expression over the whole hippocampal formation, while greater time spent in outdoor areas (the grassy range and stone yard) was associated with higher proliferation in the rostral subregion. Basal proliferation in the chicken hippocampal formation may thus relate to reactivity, while levels in the rostral region may be stimulated by ranging experience. Doublecortin expression in the caudal hippocampus negatively co-varied with time on the grassy range, but was not associated with tonic immobility duration. This suggests that ranging outside may be associated with stress. Within laying hen flocks, individual differences in hippocampal plasticity thus relate to coping style and use of external areas.

Putative adult neurogenesis in palaeognathous birds: The common ostrich (Struthio camelus) and emu (Dromaius novaehollandiae).

In the current study, we examined adult neurogenesis throughout the brain of the common ostrich (Struthio camelus) and emu (Dromaius novaehollandiae) using immunohistochemistry for the endogenous markers PCNA which labels proliferating cells, and DCX, which stains immature and migrating neurons. The distribution of PCNA and DCX labelled cells was widespread throughout the brain of both species. The highest density of cells immunoreactive to both markers was observed in the olfactory bulbs and the telencephalon, especially the subventricular zone of the lateral ventricle. Proliferative hot spots, identified with strong PCNA and DCX immunolabelling, were identified in the dorsal and ventral poles of the rostral aspects of the lateral ventricles. The density of PCNA immunoreactive cells was less in the telencephalon of the emu compared to the common ostrich. Substantial numbers of PCNA immunoreactive cells were observed in the diencephalon and brainstem, but DCX immunoreactivity was weaker in these regions, preferentially staining axons and dendrites over cell bodies, except in the medial regions of the hypothalamus where distinct DCX immunoreactive cells and fibres were observed. PCNA and DCX immunoreactive cells were readily observed in moderate density in the cortical layers of the cerebellum of both species. The distribution of putative proliferating cells and immature neurons in the brain of the common ostrich and the emu is widespread, far more so than in mammals, and compares with the neognathous birds, and suggests that brain plasticity and neuronal turnover is an important aspect of cognitive brain functions in these birds.

Generating brain matrices for zebra finch brain sectioning using three-dimensional printing technology.

BACKGROUND: The demand to sample brain regions in non-model species is increasing as more studies are integrating neurological data into behavioural, ecological or evolutionary analysis. However, the sampling operation is difficult for researchers without neuroscience background. It is also a challenge to collect neuroanatomical regions from animals in the field. NEW METHOD: Here we developed a new brain matrix for guiding researchers to section zebra finches' (Taeniopygia guttata) brains more steadily than by freehand trimming. Based on the 3D printing technology, we produced the zebra finch brain matrix from scratch. We also provided a step-by-step protocol to make brain matrices for any species with a brain size between that of shrews and dogs. RESULTS: The brain matrix could guide us to find the zebra finch's neuroanatomical landmarks, such as the hypothalamus, optic chiasm and occulomotor nerve. The matrix's channels near these landmarks could be used to section brains steadily and rapidly. COMPARISON WITH EXISTING METHODS: Standardized brain sectioning often requires expensive machines that may not be available in most laboratories or in the field, such as microtomes. In addition, machine-based trimming is time-consuming. Although commercial brain matrices can overcome these problems, they are only available for rats and mice. The brain matrices we developed are affordable to most laboratories and can be customised for non-model species in both lab and field experiments. CONCLUSIONS: The matrix-guided approach requires a relatively short training period and can allow researchers to properly and quickly sample brains, and thus will facilitate neuroscience-based interdisciplinary research.

Neurophysiology of Avian Sleep: Comparing Natural Sleep and Isoflurane Anesthesia.

Propagating slow-waves in electroencephalogram (EEG) or local field potential (LFP) recordings occur during non-rapid eye-movement (NREM) sleep in both mammals and birds. Moreover, in both, input from the thalamus is thought to contribute to the genesis of NREM sleep slow-waves. Interestingly, the general features of slow-waves are also found under isoflurane anesthesia. However, it is unclear to what extent these slow-waves reflect the same processes as those giving rise to NREM sleep slow-waves. Similar slow-wave spatio-temporal properties during NREM sleep and isoflurane anesthesia would suggest that both types of slow-waves are based on related processes. We used a 32-channel silicon probe connected to a transmitter to make intra-cortical recordings of the visual hyperpallium in naturally sleeping and isoflurane anesthetized pigeons (Columba livia) using a within-bird design. Under anesthesia, the amplitude of LFP slow-waves was higher when compared to NREM sleep. Spectral power density across all frequencies (1.5-100 Hz) was also elevated. In addition, slow-wave coherence between electrode sites was higher under anesthesia, indicating higher synchrony when compared to NREM sleep. Nonetheless, the spatial distribution of slow-waves under anesthesia was more comparable to NREM sleep than to wake or REM sleep. Similar to NREM sleep, slow-wave propagation under anesthesia mainly occurred in the thalamic input layers of the hyperpallium, regions which also showed the greatest slow-wave power during both recording conditions. This suggests that the thalamus could be involved in the genesis of slow-waves under both conditions. Taken together, although slow-waves under isoflurane anesthesia are stronger, they share spatio-temporal activity characteristics with slow-waves during NREM sleep.

Expression of aggressiveness modulates mesencephalic c-fos activation during a social interaction test in Japanese quail (Coturnix japonica).

It is well known that during a social conflict, interactions are dependent on the animal's propensity to behave aggressively as well as the behavior of the opponent. However, discriminating between these two confounding factors was difficult. Recently, a Social Interaction (SI) test using photocastrated males as non-aggressive stimuli was proposed as a useful tool to evaluate aggressiveness. The avian Intercollicular- Griseum centralis complex (comparable to mammalian periaqueductal gray) has been reported as a crucial node in the descending pathways that organize behavioral and autonomic aspects of defensive responses and aggressiveness. Herein, using the SI test, we evaluated whether mesencephalic areas are activated (expressed c-fos) when photostimulated adult males are confronted with non-responsive (non-aggressive) opponents. Furthermore, we also examined whether mesencephalic activation is related to male performance during the SI test (i.e., aggressive vs. non-aggressive males) in birds reared in enriched or in standard environments. Five mesencephalic areas at two anatomic levels (intermediate and rostral) and locomotion during SI testing were studied. Aggressive males showed increased c-fos expression in all areas studied, and moved at faster speeds in comparison to their non-aggressive and control counterparts. Non-aggressive males and the test controls showed similar c-fos labeling. In general, rearing condition did not appear to influence c-fos expression nor behavior during the SI test. Findings suggest that mesencephalic activation is involved when males are actively expressing aggressive behaviors. This overall phenomenon is shown regardless of both the environmental stimuli provided during the birds´ rearing and the potentially stressful stimuli during the SI trial.

Brain Distribution and Sexually Dimorphic Expression of Amylin in Different Reproductive Stages of the Zebra Finch (Taeniopygia guttata) Suggest Roles of the Neuropeptide in Song Learning and Social Behaviour.

The expression of the recently identified neuropeptide, amylin, is restricted in rodents to the postpartum preoptic area and may play a role in the control of parental behaviours and food intake. These processes are substantially different between bird and rodent parents as birds do not lactate but often show biparental care of the offspring. To establish the presence and role of amylin in the bird brain, in the present study, we investigated the distribution of amylin in brains of adult male and female zebra finches in three different reproductive stages (i.e. paired without young, incubating eggs or provisioning nestlings) and in unpaired control birds living in same sex flocks. Amylin mRNA was identified in the hypothalamus of zebra finch by RT-PCR, which was also used to produce probes for in situ hybridisation. Subsequently, in situ hybridisation histochemistry was performed in brain sections, and the labelling signal was quantified and compared between the groups. Amylin showed a much wider brain distribution than that of rodents. A strong and, in some regions, sexually dimorphic label was found in the striatum and several brain regions of the social behavioural network in both males and females. Many regions responsible for the learning of birdsong also contained amylin-positive neurons, and some regions showed sex differences reflecting the fact that vocalisation is sexually dimorphic in the zebra finch: only males sing. Area X (Ar.X), a striatal song centre present only in males, was labelled in paired but not unpaired male. Ar.X, another song centre, the lateral part of the magnocellular nucleus of the anterior nidopallium (lMAN) also contained amylin and had higher amylin label in paired, as opposed to unpaired birds. The wider distribution of amylin in birds as compared to rodents suggests a more general role of amylin in social or other behaviours in avian species than in mammals. Alternatively, parental care in birds may be a more complex behavioural trait involving a wider set of brain regions. The sex differences in song centres, and the changes with reproductive status suggest a participation of amylin in social behaviours and related changes in the singing of males.

Crosstalk between Estrogen Withdrawal and NFκB Signaling following Penetrating Brain Injury.

OBJECTIVES: Characterized by neuroinflammation, traumatic brain injury (TBI) induces neuropathological changes and cognitive deficits. Estrogens are neuroprotective by increasing cell survival and this increase is mediated by a decrease in neuroinflammation. To further explore the relationship between estrogens, brain injury, and neuroinflammation, we examined the expression of the IKK/NFκB complex. The IKK/NFκB complex is a pleiotropic regulator of many cellular signaling pathways linked to inflammation, as well as three major cytokines (IL-1ß, IL-6, and TNF-α). We hypothesized that NFκB expression would be upregulated following injury and that this increase would be exacerbated when circulating estrogens were decreased with fadrozole (aromatase inhibitor). METHODS: Using adult zebra finches, we first determined the expression of major components of the NFκB complex (NFκB, IκB-α, and IκB-ß) following injury using qPCR. Next, male and female finches were collected at 2 time points (2 or 24 h after injury) and brain tissue was analyzed to determine whether NFκB expression was differentially expressed in males and females at either time point. Finally, we examined how the expression of NFκB changed when estrogen levels were decreased immediately after injury. RESULTS: Our study documented an increase in the expression of the major components of the NFκB complex (NFκB, IκB-α, and IκB-ß) following injury. Decreasing estrogen levels resulted in a surprising decrease in the NFκB complex studied here. DISCUSSION: These data further expand the model of how estrogens and other steroid hormones interact with the inflammatory pathways following injury and may prove beneficial when developing therapies for treatment of TBI.

Age-related and function-dependent regional alterations of free L- and D-aspartate in postembryonic chick brain.

D-aspartate (D-Asp) modulates adult neural plasticity and embryonic brain development by promoting cell proliferation, survival and differentiation. Here, developmental changes of the excitatory amino acids (EAAs) L-Glu, L-Asp and D-Asp were determined during the first postembryonic days, a time window for early learning, in selected brain regions of domestic chickens after chiral separation and capillary electrophoresis. Extracellular concentration (ECC) of EAAs was measured in microdialysis samples from freely moving chicks. ECC of D-Asp (but not L-EAAs) decreased during the first week of age, with no considerable regional or learning-related variation. ECC of L-Asp and L-Glu (but not of D-Asp) were elevated in the mSt/Ac in response to a rewarding stimulus, suggesting importance of Asp-Glu co-release in synaptic plasticity of basal ganglia. Potassium-evoked release of D-Asp, with a protracted transient, was also demonstrated. D-Asp constitutes greater percentage of total aspartate in the extracellular space than in whole tissue extracts, thus the bulk of D-Asp detected in tissue appears in the extracellular space. Conversely, only a fraction of tissue L-EAAs can be detected in extracellular space. The lack of changes in tissue D-Asp following avoidance learning indicates a tonic, rather than phasic, mechanism in the neuromodulatory action of this amino acid.

Are There Place Cells in the Avian Hippocampus?

Birds possess a hippocampus that serves many of the same spatial and mnemonic functions as the mammalian hippocampus but achieves these outcomes with a dramatically different neuroanatomical organization. The properties of spatially responsive neurons in birds and mammals are also different. Much of the contemporary interest in the role of the mammalian hippocampus in spatial representation dates to the discovery of place cells in the rat hippocampus. Since that time, cells that respond to head direction and cells that encode a grid-like representation of space have been described in the rat brain. Research with homing pigeons has discovered hippocampal cells, including location cells, path cells, and pattern cells, that share some but not all properties of spatially responsive neurons in the rodent brain. We have recently used patterns of immediate-early gene expression, visualized by the catFISH method, to investigate how neurons in the hippocampus of brood-parasitic brown-headed cowbirds respond to spatial context. We have found cells that discriminate between different spatial environments and are re-activated when the same spatial environment is re-experienced. Given the differences in habitat and behaviour between birds and rodents, it is not surprising that spatially responsive cells in their hippocampus and other brain regions differ. The enormous diversity of avian habitats and behaviour offers the potential for understanding the general principles of neuronal representation of space.

Putative adult neurogenesis in two domestic pigeon breeds (Columba livia domestica): racing homer versus utility carneau pigeons.

Generation of neurons in the brains of adult birds has been studied extensively in the telencephalon of song birds and few studies are reported on the distribution of PCNA and DCX in the telencephalon of adult non-song learning birds. We report here on adult neurogenesis throughout the brains of two breeds of adult domestic pigeons (Columba livia domestica), the racing homer and utility carneau using endogenous immunohistochemical markers proliferating cell nuclear antigen (PCNA) for proliferating cells and doublecortin (DCX) for immature and migrating neurons. The distribution of PCNA and DCX immunoreactivity was very similar in both pigeon breeds with only a few minor differences. In both pigeons, PCNA and DCX immunoreactivity was observed in the olfactory bulbs, walls of the lateral ventricle, telencephalic subdivisions of the pallium and subpallium, diencephalon, mesencephalon and cerebellum. Generally, the olfactory bulbs and telencephalon had more PCNA and DCX cells than other regions. Two proliferative hotspots were evident in the dorsal and ventral poles of the lateral ventricles. PCNA- and DCX-immunoreactive cells migrated radially from the walls of the lateral ventricle into the parenchyma. In most telencephalic regions, the density of PCNA- and DCX-immunoreactive cells increased from rostral to caudal, except in the mesopallium where the density decreased from rostral to middle levels and then increased caudally. DCX immunoreactivity was more intense in fibres than in cell bodies and DCX-immunoreactive cells included small granular cells, fusiform bipolar cells, large round and or polygonal multipolar cells. The similarity in the distribution of proliferating cells and new neurons in the telencephalon of the two breeds of pigeons may suggest that adult neurogenesis is a conserved trait as an ecological adaptation irrespective of body size.