Estradiol-17ß Is Influenced by Age, Housing System, and Laying Performance in Genetically Divergent Laying Hens (Gallus gallus f.d.).

The estrogen estradiol-17ß is known as one of the major gonadal steroid hormones with different functions in reproduction. In this study we analyzed estradiol-17ß concentration in laying hens of four pure bred chicken laying lines at four different time intervals of the laying period (17th-19th week of age, 33rd-35th week of age, 49th-51st week of age, and 72nd week of age). The high performing white egg (WLA) and brown egg (BLA) layer lines as well as the low performing white (R11) and brown (L68) layer lines were kept in both single cages and a floor housing system. We investigated whether there were differences in estradiol -17ß concentrations between lines at different ages that could be related to selection for high egg production or phylogenetic origin of the animals, and whether there was an influence of housing conditions on estradiol-17ß. Estradiol-17ß concentrations differed between high and low performing layer lines at all time intervals studied. High performing hens showed higher estradiol-17ß concentrations compared to low performing hens. In all lines, highest estradiol-17ß concentration was measured at their 49th to their 51st week of age, whereas the peak of laying intensity was observed at their 33rd to their 35th week of age. Additionally, hens with fewer opportunities for activity housed in cages showed higher estradiol-17ß concentrations than hens kept in a floor housing system with more movement possibilities. We could show that laying performance is strongly linked with estradiol -17ß concentration. This concentration changes during laying period and is also influenced by the housing system.

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.

Smaller brains in laying hens: New insights into the influence of pure breeding and housing conditions on brain size and brain composition.

During domestication, many different chicken breeds have been developed that show many alterations compared with their wild ancestors and large variability in parameters such as body size, coloring, behavior, and even brain morphology. Among the breeds, one can differentiate between commercial and noncommercial strains, and commercial strains do not usually show variability as high as noncommercial breeds but exhibit a high production rate of eggs (or meat). The breeding of high-performing laying hens, including the housing conditions of hens, is often a focus of concern for animal welfare, and to date, little is known about the correlation between housing conditions and artificial selection on brain structure. Based on an allometric approach, we compared the relative brain sizes of 2 inbred strains of laying hens (WLA and R11) with those of 7 other noncommercial chicken breeds. In addition, we examined the brain composition of laying hens and analyzed the relative sizes of the telencephalon, hippocampus, tectum opticum, and cerebellum. Half of WLA and R11 lines were kept in floor-housing systems, and the other half were kept in a single cage-housing system. Both strains of laying hens showed significantly smaller brains than the other chicken breeds. In addition, there was a significant difference between WLA and R11 hens, with R11 hens having larger brains. There was no difference in the relative brain sizes of floor-housed and cage-housed hens. WLA and R11 hens did not differ in their brain composition, but floor-housed hens showed a significantly larger cerebellum than cage-housed hens. Apparently, pure breeding over a long time and strong artificial selection for a high production of eggs is accompanied by (unintentional) selection for smaller brains. Further studies may also reveal differences in brain composition and the influence of housing conditions on brain composition.

The Effects of Domestication on the Brain and Behavior of the Chicken in the Light of Evolution.

The avian class is characterized by particularly strong variability in their domesticated species. With more than 250 breeds and highly efficient commercial lines, domestic chickens represent the outcome of a really long period of artificial selection. One characteristic of domestication is the alterations in brain size and brain composition. The influence of domestication on brain morphology has been reviewed in the past, but mostly with a focus on mammals. Studies on avian species have seldom been taken into account. In this review, we would like to give an overview about the changes and variations in (brain) morphology and behavior in the domestic chicken, taking into consideration the constraints of evolutionary theory and the sense or nonsense of excessive artificial selection.

The hippocampus of birds in a view of evolutionary connectomics.

The avian brain displays a different brain architecture compared to mammals. This has led the first pioneers of comparative neuroanatomy to wrong conclusions about bird brain evolution by assuming that the avian telencephalon is a hypertrophied striatum. Based on growing evidence from divers analysis demonstrating that most of the avian forebrain is pallial in nature, this view has substantially changed during the past decades. Further, birds show cognitive abilities comparable to or even exceeding those of some mammals, even without a "six-layered" cortex. Beside higher associative regions, most of these cognitive functions include the processing of information in the hippocampal formation (HF) that shares a homologue structure in birds and mammals. Here we show with 3D polarized light imaging (3D-PLI) that the HF of pigeons like the mammalian HF shows regional specializations along the anterior-posterior axis in connectivity. In addition, different levels of adult neurogenesis were observed in the subdivisions of the HF per se and in the most caudal regions pointing towards a functional specialization along the anterior-posterior axis. Taken together our results point to species specific morphologies but still conserved hippocampal characteristics of connectivity, cells and adult neurogenesis if compared to the mammalian situation. Here our data provides new aspects for the ongoing discussion on hippocampal evolution and mind.

The orientation of homing pigeons (Columba livia f.d.) with and without navigational experience in a two-dimensional environment.

Homing pigeons are known for their excellent homing ability, and their brains seem to be functionally adapted to homing. It is known that pigeons with navigational experience show a larger hippocampus and also a more lateralised brain than pigeons without navigational experience. So we hypothesized that experience may have an influence also on orientation ability. We examined two groups of pigeons (11 with navigational experience and 17 without) in a standard operant chamber with a touch screen monitor showing a 2-D schematic of a rectangular environment (as "geometric" information) and one uniquely shaped and colored feature in each corner (as "landmark" information). Pigeons were trained first for pecking on one of these features and then we examined their ability to encode geometric and landmark information in four tests by modifying the rectangular environment. All tests were done under binocular and monocular viewing to test hemispheric dominance. The number of pecks was counted for analysis. Results show that generally both groups orientate on the basis of landmarks and the geometry of environment, but landmark information was preferred. Pigeons with navigational experience did not perform better on the tests but showed a better conjunction of the different kinds of information. Significant differences between monocular and binocular viewing were detected particularly in pigeons without navigational experience on two tests with reduced information. Our data suggest that the conjunction of geometric and landmark information might be integrated after processing separately in each hemisphere and that this process is influenced by experience.

The Influence of Social Parameters on the Homing Behavior of Pigeons.

Homing pigeons develop preferred routes when released alone several times from the same site, but they sometimes diverge from their preferred route when subsequently released with another pigeon. Additionally, group flights show a better homing performance than solo flights. But this knowledge is based on studies involving both sexes and lacks analyses of social parameters such as mating or breeding status, even though it is known that such parameters have an influence on behavior and on motivation for specific behavioral patterns. GPS trackers were used to track 24 homing pigeons (9 breeding pairs and 6 unmated females) as they performed a familiar 10km route in various pair and group combinations. Comparisons of efficiency indices (quotient between straight-line distance and pigeon's track) reveal that unmated females show the best efficiency in single flights. Generally, group flights show the best efficiency followed by pair flights with a social partner of the opposite sex. Pair flights with the mated partner exhibit the poorest performance. Additionally, just before squabs hatching, females show a higher efficiency index when released at 8 am, compared to releases at 2 pm. Our results indicate that homing flight efficiency can provide insight into individual motivation and that social parameters have an influence on homing performance on a familiar route.

Treatment with a neem seed extract (MiteStop®) of beetle larvae parasitizing the plumage of poultry.

Beetles of the species Alphitobius diaperinus, Dermestes bicolor, and Dermestes lardarius may transmit severe agents of diseases on poultry and may in addition harm as larvae the skin and feathers thus leading to severe economic losses. The present study deals with a control measurement using a neem seed extract (MiteStop®) being diluted with tap water. It was shown that spraying of a 1:33 dilution kills both larvae and adults of these part-time parasites as was previously shown for other parasites such as mites, ticks, and blood sucking or biting insects.

Observations on effects of a neem seed extract (MiteStop®) on biting lice (mallophages) and bloodsucking insects parasitizing horses.

The hair of 300 horses belonging to short hair and long hair races had been routinely treated during the last 3 years with a neem seed extract (MiteStop®) in order to kill mallophages (e.g., specimens of the genus Werneckiella). It was found that in all cases, a hidden infestation with these biting lice had existed, which became visible when the product (diluted 1:20 with tap water) was brushed onto the hair. The mallophages left the body surface and became visible as a fine "wooly looking" layer at the tips of the hair. Furthermore, this treatment stopped the forming of dandruff of the skin of the horses, which, in case of heavy mallophage infestations, had looked like being powdered. Another interesting result of the treatment was reported by the riders. They described that the product had a considerable repellent effect on bloodsucking tabanids, mosquitoes, ceratopogonids, simuliids, as well as on licking flies. This repellency effect was noted to last for up to 7 days if the horses were not washed.

Biting and bloodsucking lice of dogs--treatment by means of a neem seed extract (MiteStop®, Wash Away Dog).

Dogs infested with lice belonging either to the group of Mallophaga (hairlings, i.e., Trichodectes canis) or Anoplura (bloodsucking lice, e.g., Linognathus setosus) were washed with the neem seed preparations MiteStop® or Wash Away Dog. It was found that a single treatment with one of these products killed both motile stages and those developing inside eggs (nits) being glued at the hair. In both cases the product had been left for 20 min onto the hair before it was washed away just with normal tap water.

Effects of a neem seed extract (MiteStop®) on mallophages (featherlings) of chicken: in vivo and in vitro studies.

Mallophages of birds (featherlings) are mostly very tiny and can even as adults better be recognized by their movements than by their elongate body shape when using just the naked eye. Since some species (e.g., the "shaft louse" Menopon gallinae, the elongate feather louse Lipeurus caponis, or Columbicola sp.) may pierce the pulp of feathers or the skin by their biting or scratching mandibles and thus lick the excreted blood, they may be extremely dangerous especially to young birds, even if they only feed by nibbling along the feather surface and/or eat epidermal debris. The present paper reports on the successful treatment of different races of fowls being severely infested with both above cited species. This in vivo treatment was done either by a short dipping of the whole fowl into the 1:33 dilution (with tap water) of a neem seed extract (MiteStop®) or by spraying them with the freshly diluted product. It was seen that the dead mallophages dropped down from the feathers as soon as they were dry again. As a precaution, a second treatment was done by some owners 1 week after the first one in order to eliminate all stages, which eventually might have hatched from untouched nits during the time interval between the two treatments. When controlling the treated fowls 4 weeks after the treatment, in no case (treated once or twice), living motile stages were diagnosed indicating the high efficacy of this nontoxic neem seed extract. When treating in vitro cutoff feathers contaminated with L. caponis, it was seen under the stereomicroscope, that the mallophages tried to run away from the 1:33 water-diluted active compound indicating that there is also a repellent effect. Treated L. caponis stopped leg movements within 3 min and died on their feathers within 1-20 min. Then, the last slight trembling movements of their legs and convulsions of their intestine stopped finally.

Asymmetry of different brain structures in homing pigeons with and without navigational experience.

Homing pigeons (Columba livia f.d.) are well-known for their homing abilities, and their brains seem to be functionally adapted to homing as exemplified, e.g. by their larger hippocampi and olfactory bulbs. Their hippocampus size is influenced by navigational experience, and, as in other birds, functional specialisation of the left and right hemispheres ('lateralisation') occurs in homing pigeons. To show in what way lateralisation is reflected in brain structure volume, and whether some lateralisation or asymmetry in homing pigeons is caused by experience, we compared brains of homing pigeons with and without navigational experience referring to this. Fourteen homing pigeons were raised under identical constraints. After fledging, seven of them were allowed to fly around the loft and participated successfully in races. The other seven stayed permanently in the loft and thus did not share the navigational experiences of the first group. After reaching sexual maturity, all individuals were killed and morphometric analyses were carried out to measure the volumes of five basic brain parts and eight telencephalic brain parts. Measurements of telencephalic brain parts and optic tectum were done separately for the left and right hemispheres. The comparison of left/right quotients of both groups reveal that pigeons with navigational experience show a smaller left mesopallium in comparison with the right mesopallium and pigeons without navigational experience a larger left mesopallium in comparison with the right one. Additionally, there are significant differences between left and right brain subdivisions within the two pigeon groups, namely a larger left hyperpallium apicale in both pigeon groups and a larger right nidopallium, left hippocampus and right optic tectum in pigeons with navigational experience. Pigeons without navigational experience did not show more significant differences between their left and right brain subdivisions. The results of our study confirm that the brain of homing pigeons is an example for mosaic evolution and indicates that lateralisation is correlated with individual life history (experience) and not exclusively based on heritable traits.

Tool-making New Caledonian crows have large associative brain areas.

Animals with a high rate of innovative and associative-based behavior usually have large brains. New Caledonian (NC) crows stand out due to their tool manufacture, their generalized problem-solving abilities and an extremely high degree of encephalization. It is generally assumed that this increased brain size is due to the ability to process, associate and memorize diverse stimuli, thereby enhancing the propensity to invent new and complex behaviors in adaptive ways. However, this premise lacks firm empirical support since encephalization could also result from an increase of only perceptual and/or motor areas. Here, we compared the brain structures of NC crows with those of carrion crows, jays and sparrows. The brains of NC crows were characterized by a relatively large mesopallium, striatopallidal complex, septum and tegmentum. These structures mostly deal with association and motor-learning. This supports the hypothesis that the evolution of innovative or complex behavior requires a brain composition that increases the ability to associate and memorize diverse stimuli in order to execute complex motor output. Since apes show a similar correlation of cerebral growth and cognitive abilities, the evolution of advanced cognitive skills appears to have evolved independently in birds and mammals but with a similar neural orchestration.

Homing pigeons as a model for the influence of experience on brain composition-including considerations on evolutionary theory.

The brain of homing pigeons seems to be functionally adapted to homing with e.g., larger hippocampi and olfactory bulbs. Furthermore, functional lateralization occurs as well in homing pigeons. Recently, the investigation of the influence of navigational experience on brain composition and lateralization revealed larger hippocampi in homing pigeons with navigational experience compared to inexperienced homing pigeons. Additionally, there are several brain structures in homing pigeons that show a volumetrical lateralization, whereas homing pigeons with navigational experience show a more lateralized brain than pigeons without navigational experience. This gives more insights in the neuronal basis of orientation and brain development in general but demonstrates as well its complexity. Plasticity and lateralization are much more correlated with individual life history than assumed up to date and have to be more considered in comparative research of evolution.

Neurobiology of the homing pigeon--a review.

Homing pigeons are well known as good homers, and the knowledge of principal parameters determining their homing behaviour and the neurological basis for this have been elucidated in the last decades. Several orientation mechanisms and parameters-sun compass, earth's magnetic field, olfactory cues, visual cues-are known to be involved in homing behaviour, whereas there are still controversial discussions about their detailed function and their importance. This paper attempts to review and summarise the present knowledge about pigeon homing by describing the known orientation mechanisms and factors, including their pros and cons. Additionally, behavioural features like motivation, experience, and track preferences are discussed. All behaviour has its origin in the brain and the neuronal basis of homing and the neuroanatomical particularities of homing pigeons are a main topic of this review. Homing pigeons have larger brains in comparison to other non-homing pigeon breeds and particularly show increased size of the hippocampus. This underlines our hypothesis that there is a relationship between hippocampus size and spatial ability. The role of the hippocampus in homing and its plasticity in response to navigational experience are discussed in support of this hypothesis.