Regulation of ipsilateral visual information within the tectofugal visual system in zebra finches.

The eyes of zebra finches are placed laterally, the foveae are looking into different directions. It is unlikely that the birds are able to process different images from both eyes simultaneously. A neural mechanism might therefore be necessary to guide the birds' attention to one of the two eyes and to reduce the processing of information of the other. Previous studies revealed that information from the ipsilateral eye is indeed suppressed on its way to the telencephalon by the activity of the contralateral eye. It has been suggested that two nuclei of the tecto-thalamic tract, nucleus subpraetectalis and nucleus interstitio praetecto subpraetectalis, are a central part of such a suppressive mechanism. Using electrophysiological recordings, we investigated the influence of these two nuclei and nucleus rotundus on the processing of binocular visual information by treating the nuclei with picrotoxin or electrolytic lesions. Deactivation of inhibitory neurons within SP/IPS leads to a significant increase of the ectostriatal responses to ipsilateral and bilateral stimulation, the responses to contralateral stimulation remain unaffected. Lesioning SP/IPS does not alter the responses to visual stimuli. Treatment of nucleus rotundus with picrotoxin increases contralaterally and bilaterally, but not ipsilaterally evoked responses. A wiring diagram is presented which interprets these findings.

Integration of information from both eyes by single neurons of nucleus rotundus, ectostriatum and lateral neostriatum in the zebra finch (Taeniopygia guttata castanotis Gould).

Although the optic nerve in birds crosses completely, visual information from the ipsilateral eye also reaches the ectostriatum, the telencephalic statibon of the tectofugal pathway, by recrossing fibers. These recrossing projections connect the contralateral tectum opticum with the ipsilateral nucleus rotundus, which in turn projects to the ectostriatum. The ectostriatum itself projects to the overlying lateral neostriatum, an area which serves an important role in sexual imprinting. This study shows that contralateral and ipsilateral information converges on single neurons within the nucleus rotundus, the ectostriatal region and the lateral neostriatum. In the three brain areas almost all isolated neurons exhibited responses to contralateral as well as bilateral visual stimuli. The number of neurons responding to ipsilateral stimuli increases from nucleus rotundus to the lateral neostriatum. We did not find any neurons driven exclusively by ipsilateral stimuli. The strength of ipsilateral responses is rather weak within the nucleus rotundus and ectostriatum, but shows a sharp increase in the lateral neostriatum. For most neurons of nucleus rotundus and ectostriatum, an additional ipsilateral stimulus did not significantly affect the response to a contralateral one. In contrast, a strong excitatory effect can be found in the neostriatum. The results are in agreement with previous evoked-potential studies and give new insights on the integration of ipsilateral and contralateral stimuli in zebra finch tectofugal visual pathway.

Neurons with complex receptive fields in the stratum griseum centrale of the zebra finch (Taeniopygia guffata castanotis Gould) optic tectum.

The electrophysiological and morphological features of visually driven neurons of the stratum griseum centrale of the zebra finch optic tectum were studied by extracellular recording and staining techniques. Stratum griseum centrale neuron responses are sustained in most cases. Receptive fields are big, up to 150 degrees of the visual field. The excitatory center (hot spot) varies in size from 1 degrees to 15 degrees. It can be mapped by small static stimuli, adapts slower than the surround, and has a shape comparable to the excitatory fields of upper-layer neurons. In contrast, the big surround shows responses only to small moving objects which elicit a typical pattern of alternating bursts and silent periods. Alternatively, the same stimuli elicit long-lasting bursts followed by strong adaption. Anatomically, stratum griseum centrale neurons are characterized by far reaching dendrites which terminate with "bottlebrush"-like endings in the upper retinorecipient layers. In addition, they are connected with retinorecipient structures by an interneuron located between layers 10 and 11. The role of the structure of inputs for the organization of the receptive fields is discussed.

Testosterone-dependent plasticity of avian forebrain neurons is not restricted to the song control system.

Testosterone is acting on brain areas involved in the control of sexual behaviour, for example the preoptic area and the song system. We now demonstrate that it also affects other avian brain areas, as exemplified here by measurement of spine densities. Depletion of testosterone by castration or application of cyproterone acetate leads to a decrease in spine density in secondary sensory areas like lateral neo- and hyperstriatum and hyperstriatum accessorium and dorsale, or in associative areas such as the caudal archi- and neostriatum. We conclude that testosterone is acting directly on the spines, and suggest that the mechanism of spine density control by hormones may have arisen because of energy demands.

Evidence for the involvement of two areas of the zebra finch forebrain in sexual imprinting.

Sexual imprinting in male zebra finches is a two-step process, including an acquisition period early in life and a stabilization process normally occuring during the first courtship attempts of the male. During the acquisition period, a young male learns about its social environment. During stabilization, which can be delayed experimentally until day 100, it develops a preference for the appropriate object for courtship behavior on the basis of its previous and acute experience. Thereafter, this preference cannot be altered again. Exploring the physiological basis for imprinting, we have previously shown that the neurons of two forebrain areas (ANC and HAD) increase their spine density in the course of the stabilization process, while in two other areas (MNH and LNH) a decrease of spine density can be observed. With the present experiments, we tested the idea that the spine density decrease in MNH and LNH is the anatomical manifestation of the imprinting process. Previous behavioral experiments have shown that exposure to a nestbox after 100 days of age stabilizes the sexual preference of a zebra finch male as well as does exposure to a female. The present study shows that nestbox exposure also reduces the spine density in MNH and LNH, but has no effect on ANC and HAD. It has also been shown previously that treating males with an antiandrogen between days 40 and 100 affects the final preference of a male. The present experiment indicates that the same treatment affects spine growth during development in MNH and LNH and prevents the increase of spine density within HAD and ANC normally induced by exposure to a female. The results are interpreted as strong evidence for the involvement of MNH and LNH in sexual imprinting.

Behavioural and neurophysiological aspects of sexual imprinting in zebra finches.

Sexual imprinting has been defined as the process by which young animals learn the characteristics of their future sexual partners. It is a two stage process including an acquisition period where features of the social environment are learnt, and a stabilization process by which, under the guidance of the previously acquired social information, a preference for a sexual partner is established and stabilized, so that it cannot be altered again subsequently. The stabilization process is short (1 h) and can be controlled experimentally. This allows for the design of experiments to examine the physiological events accompanying the imprinting process. During the stabilization process, four areas of the forebrain are more activated than in any other behavioural context. These are the hyperstriatum accessorium/dorsale (HAD), the archi-neostriatum caudale (ANC), the medial neo/hyperstriatum (MNH) and the lateral neo/hyperstriatum (LNH). Isolation during development reduces the spine density of neurons in HAD and ANC and enhances it in MNH and LNH. Subsequent exposure to a female (which stabilizes the previously acquired preference in behavioural experiments) for 1 week leads to an enhancement of spine densities in HAD and ANC, and to a reduction in MNH and LNH. The enhancement in HAD and ANC is reversible by a second isolation period after the exposure to a female, the reduction within MNH and LNH is not. This irreversibility indicates that the reduction process within MNH and LNH may be the anatomical manifestation of the imprinting process. The examination of spine densities in the four brain areas after two experiments which have been shown previously to affect the stabilization process in behavioural experiments, confirms this idea.

Spine density changes in forebrain areas of the zebra finch by TEA-induced potentiation.

When young zebra finch males who have been reared in isolation court a female for the first time they develop a stable preference for females of this species (sexual imprinting). During this first courtship, two areas of the forebrain show a reversible enhancement of spine density, while in two other areas, spine density decreases irreversibly. Here we show that the same alterations can be induced by application of tetraethylammonium (TEA) to slices of adult, previously isolated males. TEA application induces an enhancement of spine density in two forebrain areas and leads to a decrease of spine density in two others. Although the exact mechanisms are unknown, our results indicate that potentiation phenomena are involved in the spine density changes induced by first courtship.

On the structure and function of the tectofugal visual pathway in laterally eyed birds.

Of the two visual systems in vertebrates, the tectofugal pathway has often been attributed to stimulus localization, while the thalamofugal pathway was thought to be involved in stimulus identification. This review provides evidence that the tectofugal pathway serves both functions. It is speculated that the initial task of the tectum is to localize an object and to guide the fixation movement. The object in focus is then analysed by the higher stations of the tectofugal system. The result of the analysis is fed into the optic tectum and is used for decision about the treatment of the object.

Activity-dependent plasticity in visual forebrain areas of the zebra finch.

It has previously been shown that the activity of some area of the forebrain of birds is dependent on the arousal level of the animal. Other areas do not show this dependency. This paper, on the basis of 2-DG experiments and spine density measurements on Golgi-impregnated tissue, shows that primary telencephalic target areas of the two visual pathways of zebra finch males are not dependent on arousal for activation. In contrast, secondary areas of both visual pathways show arousal-dependent activation. Only the secondary visual areas also show effects of rearing conditions on the spine density: isolation of the birds from day 40 reduces spine density in the hyperstriatum accessorium (HA) of the thalamofugal pathway, and enhances spine density in the lateral neo/hyperstriatum (LNH, tectofugal pathway) significantly from day 80, if compared to aviary-reared birds. A 1-week exposure to a female eliminates the isolation effects in both areas. A second isolation period again reduces the spine density in the HA, but does not enhance it again in the LNH. By comparison with previous studies, we conclude that the spine density in the HA reflects the complexity of the social environment. The irreversible reduction of spine density in the LNH as consequence of the 7-day exposure to a female is interpreted as physiological correlate of an imprinting process, which has previously been shown to occur at the same time. The effects in both area, the HA and LNH, are dependent on arousal, which may be mediated by brainstem efferents innervating the secondary, but not the primary, visual areas in birds and in mammals.

GABAergic inputs to the nucleus rotundus (pulvinar inferior) of the pigeon (columba livia).

The avian nucleus rotundus, a nucleus that appears to be homologous to the inferior/ caudal pulvinar of mammals, is the major target of an ascending retino-tecto-thalamic pathway. Further clarification of the inputs to the rotundus and their functional properties will contribute to our understanding of the fundamental role of the ascending tectal inputs to the telencephalon in all vertebrates, including mammals. We found that the rotundus contains a massive plexus of glutamic acid decarboxylase (GAD)-immunoreactive axons using antibodies against GAD. The cells within the rotundus, however, were not immunoreactive for GAD. The retrograde tracer cholera toxin B fragment was injected into the rotundus to establish the location of the afferent neurons and determine the source of the gamma-aminobutyric acid (GABA) inputs into the rotundus. In addition to the recognized bilateral inputs from layer 13 of the tectum, we found intense retrograde labeling of neurons within the ipsilateral nuclei subpretectalis (SP), subpretectalis-caudalis (SPcd), interstitio-pretecto-subpretectalis (IPS), posteroventralis thalami (PV), and reticularis superior thalami (RS). All the neurons of the SP, SPcd, IPS, and PV were intensely GAD-immunoreactive. The neurons of layer 13 of the tectum were not immunoreactive for GAD. Following the destruction of the ipsilateral SP/IPS complex, we found a major reduction in the intensity of the GAD axonal immunoreactivity within the ipsilateral rotundus, but this destruction did not diminish the intensity of the GAD-immunoreactivity within the contralateral rotundus. Our studies indicated that the source of the massive GAD-immunoreactive plexus within the rotundus was from the ipsilateral SP, SPcd, IPS, and PV nuclei. These nuclei, in turn, received ipsilateral tectal input via collaterals of the neurons of layer 13 in the course of their projections upon the rotundus. We suggest that the direct bilateral tecto-rotundal projections are excitatory, whereas the indirect ipsilateral projections from the SP/IPS and PV are mainly inhibitory, possibly acting via a GABA-A receptor.

Visual system alterations in White Zebra Finches.

Visual system anomalies in albino mammals are generally seen to be caused by a lack of retinal pigment and misrouting of retinofugal optic fibers. This study shows that the central visual system of white zebra finches is physiologically different from normally colored (wild type) birds, although the eye pigmentation and the retinofugal projection appear to be normal. Ipsilaterally evoked potentials in our white birds are enhanced in comparison to wild type birds, whereas in albino mammals the ipsilateral component of visually evoked potentials is reduced. Picrotoxin-induced blockade of inhibitory synapses in the ectostriatum reveals remarkable differences between wild type and white zebra finches. In wild type zebra finches, a significant shift of ipsilateral to contralateral stimulus response ratios is observed. However, there is no detectable shift in the white morph. The data suggest that inhibition of ipsilateral stimulus processing, as observed in wild type zebra finches, is significantly reduced in the white morph. Our results indicate that the effects observed in white zebra finches cannot be explained by the theories that have been developed for albinotic animals. We assume that in white zebra finches a genetic defect, which causes the white plumage, is coupled with the demonstrated deviations of inhibitory mechanisms in the central visual system.

Morphological alterations of the visual system in white zebra finches.

Visual system anomalies in albino mammals are generally caused by a lack of retinal pigment and misrouting of retinofugal optic fibres. This study shows that the visual system of white zebra finches differs morphologically from that of normally coloured (wild type) birds, although eye pigmentation and retinofugal projection are normal. In white zebra finches, the recrossing tectorotundal projection is enhanced and a new (reciprocal) connection is established between the n. rotundi of both hemispheres. The morphological alterations found in this study may contribute to the previously described enhanced processing of ipsilateral stimuli in white zebra finches.

Phase specific morphological changes induced by social experience in two forebrain areas of the zebra finch.

We examined the changes of spine density in Golgi preparations of two different areas of the forebrain of the zebra finch, the ANC (Archi-Neostriatum caudale) and MNH (medial Neo-Hyperstriatum) during development, after transferring male birds from isolation to a social condition (exposure to a female for 1 week), and after a second isolation period. MNH and ANC are two of four brain regions which are strongly activated if a male bird is exposed to a female after some time of isolation. The results of our study can be summarized as follows. 1: a peak-decline trend is observed in ANC, but not in MNH. 2: rearing conditions do not affect the development of both areas until day 70. 3: from 80 days of age, isolation leads to reduced spine density within ANC, but to enhanced spine density within MNH. 4: short social contact after isolation diminishes or eliminates the effects of isolation by an enhancement of spine density in ANC and a reduction of spine density within MNH. 5: the effects of short social rearing after isolation are reversible within ANC, but not within MNH. We presume that the alterations of spine density, which are induced by changes in social conditions, are restricted to ages older than 70 days by hormonal factors. We propose that the complexity of the ANC neuronal net follows the complexity of the social environment, and that the level of arousal is the most important factor influencing the complexity.(ABSTRACT TRUNCATED AT 250 WORDS)

[Development and adaptation of tensile strength by bones in the extremities in response to physical training exemplified by the tibia]. / Entwicklung und Anpassung der Biegefestigkeit des Extremitätenskelettes durch Training am Beispiel der Tibia.

Results from an in vivo assessment of the bending stiffness of human tibiae with a new method demonstrate that bone mineral measurements are not a suitable predictor to evaluate changes of mechanical properties of long bones. In a study on 559 male military recruits, the bone mineral at tibial shaft resulted in a mean increase of +1.8% during 15 weeks of exercise. The bending stiffness however increased about 25%. An additional test 24 months later on a sample indicated that the increase of bone mineral content was only due to the natural maturation of bone. The bending stiffness however, decreased by about 6% demonstrating the earlier training effect. No correlation between bone mineral and bending stiffness could be found neither in absolute values nor in difference between the three measurements. At the same time first results of a normative study on children (9 to 18 years old, male and female) and on women (up to 80 years old) are presented.

Spine morphology of neurons in the avian forebrain is affected by rearing conditions.

An area of the caudal forebrain of male zebra finches, the Archi-Neostriatum caudale (ANC), which is active during arousal (Bischof & Herrmann, 1986, 1988), shows rearing-dependent changes in neuron morphology (Rollenhagen & Bischof, 1991). We demonstrate here that rearing conditions also affect the shape of spines of one of the four ANC neuron types. This neuron type was examined in birds reared under five different conditions--in isolation (1), caged (2), in the aviary (3), and with social contact (4) or chasing (5) after an isolation period. Our results show that social experience determines the proportion of the three types of spines (thin, mushroom, and stubby) of the investigated neuron type. Rearing conditions and short social contact also affect the spine stem length of the thin spine type. Long-term isolation results in a reduction in number and elongation of shafts of thin spines, along with an increase of stubby-and mushroom-shaped spines. Short-term social contact or arousal enhances the number of mushroom-and thin-shaped spines and reduces the length of spine stems of thin spines. We suggest that isolation prevents the ANC neuron from reaching full development. The increase of mushroom and thin spine types due to social contact indicates that the stubby-shaped spines are replaced by, or transformed into, mushroom-shaped spines, and the mushroom-shaped spines are replaced by, or transformed into, thin spines. These results confirm and extend the experimental background for our hypothesis (Rollenhagen & Bischof, 1991) that social contact is necessary for development of normal morphology of ANC neurons.

Visual wulst influences on flash evoked responses in the ectostriatum of the zebra finch.

Anatomical data suggest that visual information from the thalamofugal pathway contributes to visual processing in the tectofugal pathway. We addressed the question of the functionality of anatomically described connections to the visual system of a laterally eyed bird, the zebra finch. The study shows the contribution of visual wulst efferents, to visual processing in the ectostriatum by recordings of visually evoked slow field potentials. Suppression of visual wulst activity resulted in a selective reduction of distinct potential components in contralaterally evoked slow field potentials. A clear reduction was observed in the maximum amplitude of short latency components in the negative wave. Long latency components of the negative wave and the entire positive wave of the contralaterally flash evoked potentials were almost abolished. Ipsilateral visual evoked potentials (VEPs) were not significantly affected. Cooling and spreading depression of the optic tectum resulted in a uniform amplitude reduction of the negative wave. The positive wave was almost abolished. Ipsilateral VEPs disappeared completely during suppression of optic tectum activity. The results showed that the visual wulst has a significant, most likely facilitatory, influence on the processing of contralateral visual information in the ectostriatum. Ipsilateral stimulus processing was partly independent from visual wulst activity. A model for thalamo- and tectofugal connectivity in the ectostriatum is suggested.

Rearing conditions affect neuron morphology in a telencephalic area of the zebra finch.

The morphology of ANC (Archi-neostriatum caudale) neurons in zebra finches is affected by arousal and rearing conditions. Branching index and spine density of ANC neurons are decreased in isolated birds and enhanced in cage reared animals, compared to aviary reared animals. Chasing the birds around the cage, or seven days of social contact with a female, raises these indices in birds isolated until adulthood relative to those of the aviary reared animals. We conclude that branching index and spine density of ANC neurons are determined during development by the amount of social contact, arousal, and activation of ANC. The changes observed after short term treatments in adult bird may depend on the same factors.

Development of flash-evoked responses in the ectostriatum of the zebra finch: an evoked potential and current-source-density analysis.

The morphological development of the tectofugal pathway in the zebra finch has recently been described in a series of studies from our laboratory. No data are currently available on the development of visual responsiveness in this pathway. We therefore investigated the development of visually evoked potentials (VEPs) in the ectostriatum, the telencephalic target area of the tectofugal pathway. Contralateral VEPs could already be recorded in 20-day-old birds, whereas ipsilateral VEPs could first be recorded in 40-day-old birds. The latencies of contralateral VEPs decrease to adult values between 20 and 40 days of age, probably due to an increase in the myelination of afferent fibers. The amplitudes of the contralateral VEPs increase continuously from day 20 to day 60; however, between 60 and 80 days of age the responses diminish substantially (-60%). Thus, contralateral VEPs in 80-day-old birds are not significantly different from those in 20-day-old birds. Thereafter the responses recover and reach their final amplitude values at about 150 days of age. The relationship of these results to morphological studies and possible mechanisms which may cause the double-peaked development of visual-evoked potentials in the ectostriatum are discussed.

Differences between ipsilaterally and contralaterally evoked potentials in the visual wulst of the zebra finch.

The telencephalic target of the thalamofugal visual pathway in birds, the visual wulst, is part of the hyperstriatum accessorium/dorsale in the bird's brain. In this study, we tried to determine the exact location of the visually responsive area in the zebra finch by recording visually evoked potentials (VEPs) from different sites throughout the hyperstriatum and calculating current source densities (CSDs). In addition, we examined the influence of ipsilateral and contralateral stimuli on stimulus processing within this area, and tried to get insight into the neuronal machinery of the thalamofugal pathway by application of drugs such as tetrodotoxin (TTX) and picrotoxin. About two-thirds of the hyperstriatum is responsive to contralateral stimuli but only a small portion responds to ipsilateral stimuli. Contralateral visual information arrives in the hyperstriatum dorsale (HD) and is processed further to the hyperstriatum accessorium (HA). The small influence of ipsilaterally evoked potentials is not due to inhibition by the activity of the contralateral eye, as could be demonstrated previously for the ectostriatum. Instead, our results show that ipsilaterally evoked potentials are inhibited at least in part by a projection from the contralateral visual wulst.

Ipsilaterally evoked responses of the zebra finch visual wulst are reduced during ontogeny.

Visual wulst responses to ipsi- and contralateral visual stimuli were investigated in young zebra finches (Taeniopygia guttata castanotis Gould) of different ages. Contralateral responses in 20, 40, 60 and 80 day old birds do not differ significantly from those in adults. In contrast, ipsilateral responses decrease substantially during development and become very weak and irregular in adult birds.