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.

Quality not quantity: Deficient juvenile play experiences lead to altered medial prefrontal cortex neurons and sociocognitive skill deficits.

Reduced play experience over the juvenile period leads to adults with impoverished social skills and to anatomical and physiological aberrations of the neurons found in the medial prefrontal cortex (mPFC). Even rearing rats from high-playing strains with low-playing strains show these developmental consequences. In the present study, we evaluated whether low-playing rats benefit from being reared with higher playing peers. To test this, we reared male Fischer 344 rats (F344), typically thought to be a low-playing strain, with a Long-Evans (LE) peer, a relatively high-playing strain. As juveniles, F344 rats reared with LE rats experienced less play and lower quality play compared to those reared with another F344. As adults, the F344 rats reared with LE partners exhibited poorer social skills and the pyramidal neurons of their mPFC had larger dendritic arbors than F344 rats reared with same-strain peers. These findings show that being reared with a more playful partner does not improve developmental outcomes of F344 rats, rather the discordance in the play styles of F344 and LE rats leads to poorer outcomes.

The relative sizes of nuclei in the oculomotor complex vary by order and behaviour in birds.

Eye movements are a critical component of visually guided behaviours, allowing organisms to scan the environment and bring stimuli of interest to regions of acuity in the retina. Although the control and modulation of eye movements by cranial nerve nuclei are highly conserved across vertebrates, species variation in visually guided behaviour and eye morphology could lead to variation in the size of oculomotor nuclei. Here, we test for differences in the size and neuron numbers of the oculomotor nuclei among birds that vary in behaviour and eye morphology. Using unbiased stereology, we measured the volumes and numbers of neurons of the oculomotor (nIII), trochlear (nIV), abducens (nVI), and Edinger-Westphal (EW) nuclei across 71 bird species and analysed these with phylogeny-informed statistics. Owls had relatively smaller nIII, nIV, nVI and EW nuclei than other birds, which reflects their limited degrees of eye movements. In contrast, nVI was relatively larger in falcons and hawks, likely reflecting how these predatory species must shift focus between the central and temporal foveae during foraging and prey capture. Unexpectedly, songbirds had an enlarged EW and relatively more nVI neurons, which might reflect accommodation and horizontal eye movements. Finally, the one merganser we measured also has an enlarged EW, which is associated with the high accommodative power needed for pursuit diving. Overall, these differences reflect species and clade level variation in behaviour, but more data are needed on eye movements in birds across species to better understand the relationships among behaviour, retinal anatomy, and brain anatomy.

Endocast structures are reliable proxies for the sizes of corresponding regions of the brain in extant birds.

Endocasts are increasingly relied upon to examine avian brain evolution because they can be used across extant and extinct species. The endocasts of birds appear to be relatively faithful representatives of the external morphology of their brains, but it is unclear how well the size of a surface feature visible on endocasts reflects the volume of the underlying brain region. The optic lobe and the Wulst are two endocast structures that are clearly visible on the external surface of avian endocasts. As they overlie two major visual regions of the brain, the optic tectum and hyperpallium, the surface areas of the optic lobe and Wulst, respectively, are often used to infer visual abilities. To determine whether the surface area of these features reflects the volume of the underlying brain regions, we compared the surface areas of the optic lobes and Wulsts from digital endocasts with the volumes of the optic tecta and hyperpallia from the literature or measured from histological series of brains of the same species. Regression analyses revealed strong, statistically significant correlations between the volumes of the brain regions and the surface areas of the overlying endocast structures. In other words, the size of the hyperpallium and optic tectum can be reliably inferred from the surface areas of the Wulst and optic lobe, respectively. This validation opens the possibility of estimating brain-region volumes for extinct species in order to gain better insights in their visual ecology. It also emphasizes the importance of adopting a quantitative approach to the analysis of endocasts in the study of brain evolution.

Selection for Divergent Reproductive Investment Affects Neuron Size and Foliation in the Cerebellum.

The cerebellum has a highly conserved internal circuitry, but varies greatly in size and morphology within and across species. Despite this variation, the underlying volumetric changes among the layers of the cerebellar cortex or their association with Purkinje cell numbers and sizes is poorly understood. Here, we examine intraspecific scaling relationships and variation in the quantitative neuroanatomy of the cerebellum in Japanese quail (Coturnix japonica) selected for high or low reproductive investment. As predicted by the circuitry of the cerebellum, the volumes of the constituent layers of the cerebellar cortex were strongly and positively correlated with one another and with total cerebellar volume. The number of Purkinje cells also significantly and positively co-varied with total cerebellar volume and the molecular layer, but not the granule cell layer or white matter volumes. Purkinje cell size and cerebellar foliation did not significantly covary with any cerebellar measures, but differed significantly between the selection lines. Males and females from the high-investment lines had smaller Purkinje cells than males and females from the low-investment lines and males from the high-investment lines had less folded cerebella than quail from the low-investment lines. These results suggest that within species, the layers of the cerebellum increase in a coordinated fashion, but Purkinje cell size and cerebellar foliation do not increase proportionally with overall cerebellum size. In contrast, selection for differential reproductive investment affects Purkinje cell size and cerebellar foliation, but not other quantitative measures of cerebellar anatomy.

Allometric Scaling Rules of the Cerebellum in Galliform Birds.

Although the internal circuitry of the cerebellum is highly conserved across vertebrate species, the size and shape of the cerebellum varies considerably. Recent comparative studies have examined the allometric rules between cerebellar mass and number of neurons, but data are lacking on the numbers and sizes of Purkinje and granule cells or scaling of cerebellar foliation. Here, we investigate the allometric rules that govern variation in the volumes of the layers of the cerebellum, the numbers and sizes of Purkinje cells and granule cells and the degree of the cerebellar foliation across 7 species of galliform birds. We selected Galliformes because they vary greatly in body and brain sizes. Our results show that the molecular, granule and white matter layers all increase in volume at the same rate relative to total cerebellum volume. Both numbers and sizes of Purkinje cells increased with cerebellar volume, but numbers of Purkinje cells increased at a much faster rate than size. Granule cell numbers increased with cerebellar volume, but size did not. Sizes and numbers of Purkinje cells as well as numbers of granule cells were positively correlated with the degree of cerebellar foliation, but granule cell size decreased with higher degrees of foliation. The concerted changes among the volumes of cerebellar layers likely reflects the conserved neural circuitry of the cerebellum. Also, our data indicate that the scaling of cell sizes can vary markedly across neuronal populations, suggesting that evolutionary changes in cell sizes might be more complex than what is often assumed.

Zebrin Expression in the Cerebellum of Two Crocodilian Species.

While in birds and mammals the cerebellum is a highly convoluted structure that consists of numerous transverse lobules, in most amphibians and reptiles it consists of only a single unfolded sheet. Orthogonal to the lobules, the cerebellum is comprised of sagittal zones that are revealed in the pattern of afferent inputs, the projection patterns of Purkinje cells, and Purkinje cell response properties, among other features. The expression of several molecular markers, such as aldolase C, is also parasagittally organized. Aldolase C, also known as zebrin II (ZII), is a glycolytic enzyme expressed in the cerebellar Purkinje cells of the vertebrate cerebellum. In birds, mammals, and some lizards (Ctenophoresspp.), ZII is expressed in a heterogenous fashion of alternating sagittal bands of high (ZII+) and low (ZII-) expression Purkinje cells. In contrast, turtles and snakes express ZII homogenously (ZII+) in their cerebella, but the pattern in crocodilians is unknown. Here, we examined the expression of ZII in two crocodilian species (Crocodylus niloticus and Alligator mississippiensis) to help determine the evolutionary origin of striped ZII expression in vertebrates. We expected crocodilians to express ZII in a striped (ZII+/ZII-) manner because of their close phylogenetic relationship to birds and their larger and more folded cerebellum compared to that of snakes and turtles. Contrary to our prediction, all Purkinje cells in the crocodilian cerebellum had a generally homogenous expression of ZII (ZII+) rather than clear ZII+/- stripes. Our results suggest that either ZII stripes were lost in three groups (snakes, turtles, and crocodilians) or ZII stripes evolved independently three times (lizards, birds, and mammals).

Zebrin II Is Expressed in Sagittal Stripes in the Cerebellum of Dragon Lizards (Ctenophorus sp.).

Aldolase C, also known as zebrin II (ZII), is a glycolytic enzyme that is expressed in cerebellar Purkinje cells of the vertebrate cerebellum. In both mammals and birds, ZII is expressed heterogeneously, such that there are sagittal stripes of Purkinje cells with high ZII expression (ZII+) alternating with stripes of Purkinje cells with little or no expression (ZII-). In contrast, in snakes and turtles, ZII is not expressed heterogeneously; rather all Purkinje cells are ZII+. Here, we examined the expression of ZII in the cerebellum of lizards to elucidate the evolutionary origins of ZII stripes in Sauropsida. We focused on the central netted dragon (Ctenophorus nuchalis) but also examined cerebellar ZII expression in 5 other dragon species (Ctenophorus spp.). In contrast to what has been observed in snakes and turtles, we found that in these lizards, ZII is heterogeneously expressed. In the posterior part of the cerebellum, on each side of the midline, there were 3 sagittal stripes consisting of Purkinje cells with high ZII expression (ZII+) alternating with 2 sagittal stripes with weaker ZII expression (ZIIw). More anteriorly, most of the Purkinje cells were ZII+, except laterally, where the Purkinje cells did not express ZII (ZII-). Finally, all Purkinje cells in the auricle (flocculus) were ZII-. Overall, the parasagittal heterogeneous expression of ZII in the cerebellum of lizards is similar to that in mammals and birds, and contrasts with the homogenous ZII+ expression seen in snakes and turtles. We suggest that a sagittal heterogeneous expression of ZII represents the ancestral condition in stem reptiles which was lost in snakes and turtles.

Seasonal Variation in Forebrain Region Sizes in Male Ruffed Grouse (Bonasa umbellus).

The song system of songbirds has provided significant insight into the underlying mechanisms and behavioural consequences of seasonal neuroplasticity. The extent to which seasonal changes in brain region volumes occur in non-songbird species has, however, remained largely untested. Here, we tested whether brain region volumes varied with season in the ruffed grouse (Bonasa umbellus), a gallinaceous bird that produces a unique wing-beating display known as 'drumming' as its primary form of courtship behaviour. Using unbiased stereology, we measured the sizes of the cerebellum, nucleus rotundus, telencephalon, mesopallium, hippocampal formation, striatopallidal complex and arcopallium across spring males, fall males and fall females. The majority of these brain regions did not vary significantly across these three groups. The two exceptions were the striatopallidal complex and arcopallium, both of which were significantly larger in spring males that are actively drumming. These seasonal changes in volume strongly implicate the striatopallidal complex and arcopallium as key structures in the production and/or modulation of the ruffed grouse drumming display and represent the first evidence of seasonal plasticity in the telencephalon underlying a non-vocal courtship behaviour. Our findings also suggest that seasonal plasticity in the striatopallidal complex and arcopallium might be a trait that is shared across many bird species and that both structures are related to the production of multiple forms of courtship and not just learned song.

Eye Morphology and Retinal Topography in Hummingbirds (Trochilidae: Aves).

Hummingbirds are a group of small, highly specialized birds that display a range of adaptations to their nectarivorous lifestyle. Vision plays a key role in hummingbird feeding and hovering behaviours, yet very little is known about the visual systems of these birds. In this study, we measured eye morphology in 5 hummingbird species. For 2 of these species, we used stereology and retinal whole mounts to study the topographic distribution of neurons in the ganglion cell layer. Eye morphology (expressed as the ratio of corneal diameter to eye transverse diameter) was similar among all 5 species and was within the range previously documented for diurnal birds. Retinal topography was similar in Amazilia tzacatl and Calypte anna. Both species had 2 specialized retinal regions of high neuron density: a central region located slightly dorso-nasal to the superior pole of the pecten, where densities reached ∼ 45,000 cells · mm(-2), and a temporal area with lower densities (38,000-39,000 cells · mm(-2)). A weak visual streak bridged the two high-density areas. A retina from Phaethornis superciliosus also had a central high-density area with a similar peak neuron density. Estimates of spatial resolving power for all 3 species were similar, at approximately 5-6 cycles · degree(-1). Retinal cross sections confirmed that the central high-density region in C. anna contains a fovea, but not the temporal area. We found no evidence of a second, less well-developed fovea located close to the temporal retina margin. The central and temporal areas of high neuron density allow for increased spatial resolution in the lateral and frontal visual fields, respectively. Increased resolution in the frontal field in particular may be important for mediating feeding behaviors such as aerial docking with flowers and catching small insects.

Zebrin II Expression in the Cerebellum of a Paleognathous Bird, the Chilean Tinamou (Nothoprocta perdicaria).

Zebrin II (ZII) is a glycolytic enzyme expressed in cerebellar Purkinje cells. In both mammals and birds, ZII is expressed heterogeneously, such that there are sagittal stripes of Purkinje cells with a high ZII expression (ZII+) alternating with stripes of Purkinje cells with little or no expression (ZII-). To date, ZII expression studies are limited to neognathous birds: pigeons (Columbiformes), chickens (Galliformes), and hummingbirds (Trochilidae). These previous studies divided the avian cerebellum into 5 transverse regions based on the pattern of ZII expression. In the lingular region (lobule I) all Purkinje cells are ZII+. In the anterior region (lobules II-V) there are 4 pairs of ZII+/- stripes. In the central region (lobules VI-VIII) all Purkinje cells are ZII+. In the posterior region (lobules VIII-IX) there are 5-7 pairs of ZII+/- stripes. Finally, in the nodular region (lobule X) all Purkinje cells are ZII+. As the pattern of ZII stripes is quite similar in these disparate species, it appears that it is highly conserved. However, it has yet to be studied in paleognathous birds, which split from the neognaths over 100 million years ago. To better understand the evolution of cerebellar compartmentation in birds, we examined ZII immunoreactivity in a paleognath, the Chilean tinamou (Nothoprocta perdicaria). In the tinamou, Purkinje cells expressed ZII heterogeneously such that there were sagittal ZII+ and ZII- stripes of Purkinje cells, and this pattern of expression was largely similar to that observed in neognathous birds. For example, all Purkinje cells in the lingular (lobule I) and nodular (lobule X) regions were ZII+, and there were 4 pairs of ZII+/- stripes in the anterior region (lobules II-V). In contrast to neognaths, however, ZII was expressed in lobules VI-VII as a series of sagittal stripes in the tinamou. Also unlike in neognaths, stripes were absent in lobule IXab, and all Purkinje cells expressed ZII in the tinamou. The differences in ZII expression between the tinamou and neognaths could reflect behavior, but the general similarity of the expression patterns across all bird species suggests that ZII stripes evolved early in the avian phylogenetic tree.

Anatomical specializations for enhanced olfactory sensitivity in kiwi, Apteryx mantelli.

The ability to function in a nocturnal and ground-dwelling niche requires a unique set of sensory specializations. The New Zealand kiwi has shifted away from vision, instead relying on auditory and tactile stimuli to function in its environment and locate prey. Behavioral evidence suggests that kiwi also rely on their sense of smell, using olfactory cues in foraging and possibly also in communication and social interactions. Anatomical studies appear to support these observations: the olfactory bulbs and tubercles have been suggested to be large in the kiwi relative to other birds, although the extent of this enlargement is poorly understood. In this study, we examine the size of the olfactory bulbs in kiwi and compare them with 55 other bird species, including emus, ostriches, rheas, tinamous, and 2 extinct species of moa (Dinornithiformes). We also examine the cytoarchitecture of the olfactory bulbs and olfactory epithelium to determine if any neural specializations beyond size are present that would increase olfactory acuity. Kiwi were a clear outlier in our analysis, with olfactory bulbs that are proportionately larger than those of any other bird in this study. Emus, close relatives of the kiwi, also had a relative enlargement of the olfactory bulbs, possibly supporting a phylogenetic link to well-developed olfaction. The olfactory bulbs in kiwi are almost in direct contact with the olfactory epithelium, which is indeed well developed and complex, with olfactory receptor cells occupying a large percentage of the epithelium. The anatomy of the kiwi olfactory system supports an enhancement for olfactory sensitivities, which is undoubtedly associated with their unique nocturnal niche.

NSF workshop report: discovering general principles of nervous system organization by comparing brain maps across species.

Efforts to understand nervous system structure and function have received new impetus from the federal Brain Research through Advancing Innovative Neurotechnologies (BRAIN) Initiative. Comparative analyses can contribute to this effort by leading to the discovery of general principles of neural circuit design, information processing, and gene-structure-function relationships that are not apparent from studies on single species. We here propose to extend the comparative approach to nervous system 'maps' comprising molecular, anatomical, and physiological data. This research will identify which neural features are likely to generalize across species, and which are unlikely to be broadly conserved. It will also suggest causal relationships between genes, development, adult anatomy, physiology, and, ultimately, behavior. These causal hypotheses can then be tested experimentally. Finally, insights from comparative research can inspire and guide technological development. To promote this research agenda, we recommend that teams of investigators coalesce around specific research questions and select a set of 'reference species' to anchor their comparative analyses. These reference species should be chosen not just for practical advantages, but also with regard for their phylogenetic position, behavioral repertoire, well-annotated genome, or other strategic reasons. We envision that the nervous systems of these reference species will be mapped in more detail than those of other species. The collected data may range from the molecular to the behavioral, depending on the research question. To integrate across levels of analysis and across species, standards for data collection, annotation, archiving, and distribution must be developed and respected. To that end, it will help to form networks or consortia of researchers and centers for science, technology, and education that focus on organized data collection, distribution, and training. These activities could be supported, at least in part, through existing mechanisms at NSF, NIH, and other agencies. It will also be important to develop new integrated software and database systems for cross-species data analyses. Multidisciplinary efforts to develop such analytical tools should be supported financially. Finally, training opportunities should be created to stimulate multidisciplinary, integrative research into brain structure, function, and evolution.

Ecomorphology of eye shape and retinal topography in waterfowl (Aves: Anseriformes: Anatidae) with different foraging modes.

Despite the large body of literature on ecomorphological adaptations to foraging in waterfowl, little attention has been paid to their sensory systems, especially vision. Here, we compare eye shape and retinal topography across 12 species representing 4 different foraging modes. Eye shape was significantly different among foraging modes, with diving and pursuit-diving species having relatively smaller corneal diameters compared to non-diving species. This may be associated with differences in ambient light intensity while foraging or an ability to tightly constrict the pupil in divers in order to facilitate underwater vision. Retinal topography was similar across all species, consisting of an oblique visual streak, a central area of peak cell density, and no discernible fovea. Because the bill faces downwards when the head is held in the normal posture in waterfowl, the visual streak will be held horizontally, allowing the horizon to be sampled with higher visual acuity. Estimates of spatial resolving power were similar among species with only the Canada goose having a higher spatial resolution. Overall, we found no evidence of ecomorphological adaptations to different foraging modes in the retinal ganglion cell layer in waterfowl. Rather, retinal topography in these birds seems to reflect the 'openness' of their habitats.

Comparative study of visual pathways in owls (Aves: Strigiformes).

Although they are usually regarded as nocturnal, owls exhibit a wide range of activity patterns, from strictly nocturnal, to crepuscular or cathemeral, to diurnal. Several studies have shown that these differences in the activity pattern are reflected in differences in eye morphology and retinal organization. Despite the evidence that differences in activity pattern among owl species are reflected in the peripheral visual system, there has been no attempt to correlate these differences with changes in the visual regions in the brain. In this study, we compare the relative size of nuclei in the main visual pathways in nine species of owl that exhibit a wide range of activity patterns. We found marked differences in the relative size of all visual structures among the species studied, both in the tectofugal and the thalamofugal pathway, as well in other retinorecipient nuclei, including the nucleus lentiformis mesencephali, the nucleus of the basal optic root and the nucleus geniculatus lateralis, pars ventralis. We show that the barn owl (Tyto alba), a species widely used in the study of the integration of visual and auditory processing, has reduced visual pathways compared to strigid owls. Our results also suggest there could be a trade-off between the relative size of visual pathways and auditory pathways, similar to that reported in mammals. Finally, our results show that although there is no relationship between activity pattern and the relative size of either the tectofugal or the thalamofugal pathway, there is a positive correlation between the relative size of both visual pathways and the relative number of cells in the retinal ganglion layer.

Brain size and morphology of the brood-parasitic and cerophagous honeyguides (Aves: Piciformes).

Honeyguides (Indicatoridae, Piciformes) are unique among birds in several respects. All subsist primarily on wax, are obligatory brood parasites and one species engages in 'guiding' behavior in which it leads human honey hunters to bees' nests. This unique life history has likely shaped the evolution of their brain size and morphology. Here, we test that hypothesis using comparative data on relative brain and brain region size of honeyguides and their relatives: woodpeckers, barbets and toucans. Honeyguides have significantly smaller relative brain volumes than all other piciform taxa. Volumetric measurements of the brain indicate that honeyguides have a significantly larger cerebellum and hippocampal formation (HF) than woodpeckers, the sister clade of the honeyguides, although the HF enlargement was not significant across all of our analyses. Cluster analyses also revealed that the overall composition of the brain and telencephalon differs greatly between honeyguides and woodpeckers. The relatively smaller brains of the honeyguides may be a consequence of brood parasitism and cerophagy ('wax eating'), both of which place energetic constraints on brain development and maintenance. The inconclusive results of our analyses of relative HF volume highlight some of the problems associated with comparative studies of the HF that require further study.

Online repositories of photographs and videos provide insights into the evolution of skilled hindlimb movements in birds.

The ability to manipulate objects with limbs has evolved repeatedly among land tetrapods. Several selective forces have been proposed to explain the emergence of forelimb manipulation, however, work has been largely restricted to mammals, which prevents the testing of evolutionary hypotheses in a comprehensive evolutionary framework. In birds, forelimbs have gained the exclusive function of flight, with grasping transferred predominantly to the beak. In some birds, the feet are also used in manipulative tasks and appear to share some features with manual grasping and prehension in mammals, but this has not been systematically investigated. Here we use large online repositories of photographs and videos to quantify foot manipulative skills across a large sample of bird species (>1000 species). Our results show that a complex interaction between niche, diet and phylogeny drive the evolution of manipulative skills with the feet in birds. Furthermore, we provide strong support for the proposition that an arboreal niche is a key element in the evolution of manipulation in land vertebrates. Our systematic comparison of foot use in birds provides a solid base for understanding morphological and neural adaptations for foot use in birds, and for studying the convergent evolution of manipulative skills in birds and mammals.

Comparative Analysis of the Temporal Impacts of Corticosterone and Simulated Production Stressors on the Metabolome of Broiler Chickens.

The impact of physiological stress on the metabolome of breast muscle, liver, kidney, and hippocampus was investigated in Ross 308 broiler chicks. Simulated on-farm stressors were compared to a corticosterone model of physiological stress. The three different stressors investigated were: (i) corticosterone at a dose of 15 mg/kg of feed; (ii) heat treatment of 36 °C and 40% RH for 8 h per day; and (iii) isolation for 1 h per day. Liver, kidney, breast muscle, and hippocampus samples were taken after 2, 4, 6, and 8 days of stress treatment, and subjected to untargeted 1H-nuclear magnetic resonance (NMR) spectroscopy-based metabolomic analysis to provide insights on how stress can modulate metabolite profiles and biomarker discovery. Many of the metabolites that were significantly altered in tissues were amino acids, with glycine and alanine showing promise as candidate biomarkers of stress. Corticosterone was shown to significantly alter alanine, aspartate, and glutamate metabolism in the liver, breast, and hippocampus, while isolation altered the same pathways, but only in the kidneys and hippocampus. Isolation also significantly altered the glycine, serine, and threonine metabolism pathway in the liver and breast, while the same pathway was significantly altered by heat in the liver, kidneys, and hippocampus. The study's findings support corticosterone as a model of stress. Moreover, a number of potential metabolite biomarkers were identified in chicken tissues, which may allow producers to effectively monitor stress and to objectively develop and evaluate on-farm mitigations, including practices that reduce stress and enhance bird health.

Interspecifc variation in eye shape and retinal topography in seven species of galliform bird (Aves: Galliformes: Phasianidae).

Eye morphology and the retinal topography of animals that live in either 'open' (e.g., grassland) or 'enclosed' (e.g., forest) terrestrial habitats show common adaptations to constraints imposed by these different habitat types. Although relationships between habitat and the visual system are well documented in most vertebrates, relatively few studies have examined this relationship in birds. Here, we compare eye shape and retinal topography across seven species from the family Phasianidae (Galliformes) that are diurnally active in either open or enclosed habitats. Species from enclosed habitats have significantly larger corneal diameters, relative to transverse diameters, than species from open habitats, which we predict serves to enhance visual sensitivity. Retinal topography, however, was similar across all seven species and consisted of a centrally positioned area centralis and a weak horizontal visual streak, with no discernible fovea. In the Japanese quail (Coturnix japonica), there was also a dorso-temporal extension of increased neuron density and, in some specimens, a putative area dorsalis. The total number of neurons in the retinal ganglion cell layer was correlated with retinal whole-mount area. Average and peak neuron densities were similar across species, with the exception of the Japanese quail, which had greater average and peak densities. Peak anatomical spatial resolving power was also similar among species, ranging from approximately 10-13 cycles/°. Overall, the pattern of retinal topography we found in phasianids is associated with ground-foraging in birds and presumably facilitates the identification of small food items on the ground as well as other visually guided behaviors, irrespective of habitat type.

Hummingbirds have a greatly enlarged hippocampal formation.

Both field and laboratory studies demonstrate that hummingbirds (Apodiformes, Trochilidae) have exceptional spatial memory. The complexity of spatial-temporal information that hummingbirds must retain and use daily is probably subserved by the hippocampal formation (HF), and therefore, hummingbirds should have a greatly expanded HF. Here, we compare the relative size of the HF in several hummingbird species with that of other birds. Our analyses reveal that the HF in hummingbirds is significantly larger, relative to telencephalic volume, than any bird examined to date. When expressed as a percentage of telencephalic volume, the hummingbird HF is two to five times larger than that of caching and non-caching songbirds, seabirds and woodpeckers. This HF expansion in hummingbirds probably underlies their ability to remember the location, distribution and nectar content of flowers, but more detailed analyses are required to determine the extent to which this arises from an expansion of HF or a decrease in size of other brain regions.