Biomedical subjects

F Nottebohm

Publications and source records attributed to F Nottebohm.

At least 37 records · Page 2Linked to original sources

PubMed1994

Cell death and neuronal recruitment in the high vocal center of adult male canaries are temporally related to changes in song.

Adult male canaries modify their song every year. Most of these changes occur during late summer and early fall, after the end of the breeding season, and in late winter, immediately before the onset of the next breeding season. The high vocal center (HVC) is an important nucleus in the brain pathway that controls this learned behavior. New neurons continue to be added to the HVC of adult male canaries, where they replace older neurons that have died. The present report describes the monthly incidence of cell death and neuronal addition in the HVC of such birds. Different groups of 1- to 2-year-old male canaries were treated with [3H]thymidine, a marker of cell birth, during each month of the year and killed 27 days later. The ratio of 3H-labeled neurons to all neurons in the HVC showed seasonal peaks and troughs. This ratio was highest in October and March. Peaks in the ratio of pycnotic (dying) HVC cells to all neurons in HVC preceded the peaks in the ratio of 3H-labeled neurons. We suggest that seasonal peaks in cell loss and neuronal recruitment in HVC are related to endocrine changes and that all three play a role in the seasonality of song modification.

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PubMed1994

The life span of new neurons in a song control nucleus of the adult canary brain depends on time of year when these cells are born.

The number of high vocal center (HVC) neurons labeled in adult male canaries by systemic injections of [3H]thymidine depended on season and survival time. This was true for HVC neurons projecting to the robust nucleus of the archistriatum and for other HVC neurons that could not be retrogradely filled from the robust nucleus of the archistriatum. Birds injected in October and killed 40 days later had twice as many labeled HVC neurons as birds injected in May and killed 40 days later. However, this difference became much larger (5 times) when the birds were allowed to survive for 4 months. Whereas more than half of the spring-born neurons disappeared between 40 days and 4 months, there was no reduction in the number of fall-born neurons present at the 4-month survival point. We infer that seasonal variables affect the life span of HVC neurons born in adulthood.

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PubMed1994

Testosterone increases the recruitment and/or survival of new high vocal center neurons in adult female canaries.

New neurons are added to the high vocal center (HVC) of adult male and female canaries. Exogenous testosterone induces a marked increase in HVC size in adult female canaries, though the mechanisms responsible for this increase remain unknown. To understand the mechanisms, we analyzed the effects of testosterone on neuronal recruitment in the female HVC. Intact adult female canaries received Silastic implants that were empty or filled with testosterone. Birds in the short-survival group received the Silastic implant, followed by a single injection of [3H]thymidine 2 days later, and were killed on the following day. Birds in the long-survival group were injected once a day for 5 days with [3H]thymidine and received the Silastic implant 20 and 40 days later. These birds were killed 60 days after the first injection of [3H]thymidine. The number of 3H-labeled ventricular zone cells above, rostral, or caudal to HVC was not affected by the hormone treatment in the short-survival birds, suggesting that testosterone did not affect neuronal production. However, the number of 3H-labeled HVC neurons that projected to robust nucleus of the archistriatum (RA) in the long-survival birds was three times greater in the hormone-treated than in the control group, though the total number of RA-projecting cells did not change significantly. Testosterone also induced an increase in the size of the HVC cells that project to RA. Thus, these experiments suggest that testosterone affects the recruitment and/or survival of newly generated RA-projecting HVC neurons but does not affect their production.

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PubMed1993

Role of a telencephalic nucleus in the delayed song learning of socially isolated zebra finches.

Male zebra finches normally learn their song from adult models during a restricted period of juvenile development. If song models are not available then, juveniles develop an isolate song which can be modified in adulthood. In this report we investigate the features of juvenile experience that underly the timing of song learning. Juvenile males raised in soundproof chambers or in visual isolation from conspecifics developed stable isolate song. However, whereas visual isolate song notes were similar to those of colony-reared males, soundproof chamber isolates included many phonologically abnormal notes in their songs. Despite having stable isolate songs, both groups copied new notes from tutors presented to them in adulthood (2.7 notes per bird for soundproof chamber isolates, 4.4 notes per bird for visual isolates). Old notes were often modified or eliminated. We infer that social interactions with live tutors are normally important for closing the sensitive period for song learning. Lesions of a forebrain nucleus (IMAN) had previously been shown to disrupt juvenile song learning, but not maintenance of adult song for up to 5 weeks after surgery. In this study, colony-reared adult males given bilateral lesions of IMAN retained all their song notes for up to 4-7.5 months after lesioning. However, similar lesions blocked all song note acquisition in adulthood by both visual and soundproof chamber isolates. Other work has shown that intact hearing is necessary for the maintenance of adult zebra finch song. We infer that auditory pathways used for song maintenance and acquisition differ: IMAN is necessary for auditorily guided song acquisition--whether by juveniles or adults--but not for adult auditorily guided song maintenance.

Aging↗

PubMed1993

Direct evidence for loss and replacement of projection neurons in adult canary brain.

Normally occurring projection neuron loss and replacement were quantified over a 6 month period in the pathway from the high vocal center (HVC) to the robust nucleus of the archistriatum (RA) in adult male canaries. Fluorescent latex microspheres were injected into RA in April--a procedure resulting in long-term retrograde labeling of RA-projecting HVC neurons. Labeled cell densities were then obtained 4 and 20 d later in April and 195 d later in October. We found that 41-49% of the RA-projecting HVC neurons present the previous April were no longer present in October. Fluorogold injections in RA 3 d prior to death in April and October retrogradely labeled similar overall densities of RA-projecting HVC neurons, indicating that cells lost over this 6 month period were replaced by new RA-projecting HVC neurons. Newer cells were larger than older cells, suggesting that an age-dependent reduction in size might precede death. Over the same time interval, no loss was observed for neurons projecting from the lateral magnocellular nucleus of the anterior neostriatum to RA. Thus, loss was specific to the input from HVC to RA. These findings raise the possibility that much if not all of the pathway from HVC to RA is replaced within a year. The time period examined encompasses the yearly transition from stable song to song learning in the canary (Nottebohm et al., 1986, 1987). A pronounced loss and replacement of neurons implicated in vocal control during this period may relate to the canary's ability to modify song in adulthood.

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PubMed1992

Role of gender, season, and familiarity in discrimination of conspecific song by zebra finches (Taeniopygia guttata).

Operant conditioning techniques were used to assess how gender and song familiarity affect song discrimination in adult zebra finches (Taeniopygia guttata). Twenty-five males and females, divided into five cohorts, were trained to discriminate between conspecific songs at different times of year. Males discriminating between their own and another song from their own aviary reached criterion in the fewest number of trials, followed by males discriminating between two songs from their own aviary, then by males discriminating between songs they had not heard before. Females discriminating between two songs from their own aviary required more trials than males to reach criterion, but, unlike in males, song familiarity did not have a significant effect on song discrimination by females. The number of trials required to reach criterion was greater in winter than in summer, suggesting a photoperiodic effect in what has been regarded as a nonphotoperiodic species. Gender, season, and familiarity appear to affect how zebra finches discriminate between conspecific songs.

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PubMed1992

Hemispheric differences in avian song discrimination.

Auditory input to the right or left forebrain of adult male zebra finches (Taeniopygia guttata) was disrupted by lesioning the ipsilateral auditory relay nucleus of the thalamus. These birds were then presented with two kinds of auditory discriminations: (i) between their own song and the song of a cage mate; (ii) between two versions of an unfamiliar zebra finch song that differed only in the harmonic profile of one of the syllables. Right-side lesion birds did better than left-side lesion ones at discriminating between their own song and the song of a cage mate; left-side lesion birds did better on the harmonic profile task. We suggest that the two halves of the zebra finch brain process conspecific sounds differently, as seems to be the case for humans.

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PubMed1992

Testosterone facilitates some conspecific song discriminations in castrated zebra finches (Taeniopygia guttata).

An experiment was designed to test for the influence of testosterone on song discriminations. We found that testosterone did have an effect, which interacted with practice and the nature of the stimuli. Fourteen adult castrated zebra finches (Taeniopygia guttata) were grouped into seven pairs. In each pair, one bird was implanted with a testosterone-filled silastic tube and the other was implanted with an empty silastic tube. They were then trained on a go/no-go operant task to discriminate between bird songs in six consecutive experiments. The songs to be discriminated were as follows: experiment 1, two canary song segments; experiment 2, the bird's own song and that of the other member of the pair; experiment 3, the same two songs as in experiment 2 but with reversed stimulus-response contingencies; experiment 4, two other zebra finch songs; experiment 5, another two zebra finch songs; and experiment 6, another two canary song segments. There were no reliable learning differences between birds treated with testosterone or with an empty silastic in experiments 1 and 3-6. However, in experiment 2, testosterone-treated birds mastered the discrimination between their own song and the song of the other member of the pair in fewer trials than birds treated with empty silastics. We suggest that a song's ability to control the behavior of male zebra finches is influenced by the nature of the song, prior experience with the training paradigm, and hormone levels.

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PubMed1992

High vocal center growth and its relation to neurogenesis, neuronal replacement and song acquisition in juvenile canaries.

It is generally thought that most circuits of the adult central nervous system (CNS) are sculpted, in part at least, by selective elimination of some of the neurons present in an initial overabundant set. In this scenario, the birth of neurons precedes the period when brain functions, such as learning, first occur. In contrast to this form of brain assembly, we describe here the delayed development of the high vocal center (HVC) and one of its efferent pathways in canaries. The retrograde tracer Fluoro-Gold (FG) was injected into one of HVC's two efferent targets, the nucleus robustus archistriatalis (RA), to define the boundaries of HVC. The HVC grows markedly between 1 and 4 months, invading neighboring territories of the caudal telencephalon. During this same period, 0.43%-0.64% of the HVC neurons present at 1 year of age are labeled per day of [3H]-thymidine injection. [3H]-Thymidine labeling is a marker of cell birth, and during the first 4 months HVC neuron number increases, probably accounting for part of the HVC growth observed. Thereafter, the number of HVC neurons remains constant, but neuronal birth persists. We infer from this that neuronal replacement starts as early as 4 months after hatching and perhaps before then. About half of the neurons born after posthatching day 10 grow an axon to RA to form the main efferent pathway exiting from HVC. HVC growth, neurogenesis, axogenesis, and the observed replacement of neurons happen during the period of juvenile vocal learning. However, the recruitment of neurons that are still present at 1 year shows no particular inflections corresponding to the various stages in song learning, and continues at essentially the same rate after the more stereotyped adult song has been acquired. We suggest that a combination of neurogenesis and neuronal replacement provides unique advantages for learning.

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PubMed1992

Right-side dominance for song control in the zebra finch.

Adult male zebra finches underwent unilateral denervation of the syrinx or unilateral lesion of the forebrain nucleus HVC known to be important for song control. Disruptive effects on song were greater after right-side than after left-side operations. After denervation of the right half of the syrinx, the fundamental frequencies of all syllables within a song converged on a value near 500 Hz, and nearly all syllables were altered in type. In contrast, the syllables produced after denervation of the left side of the syrinx largely maintained their preoperative frequencies, and fewer syllables changed in type. Unlike nerve sections, HVC lesions did not result in strikingly lateralized effects on syllable phonology; however, HVC lesions did affect the temporal patterning of a bird's song, whereas nerve sections did not, and changes in temporal patterning were more marked after right than after left HVC lesions. Right-side dominance for zebra finch song control is the reverse of that described in other songbird species with lateral asymmetry for vocal communication. We suggest that the need for a dominant side is more important than the side of dominance.

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PubMed1991

Lateral asymmetries and testosterone-induced changes in the gross morphology of the hypoglossal nucleus in adult canaries.

The caudal portion of the hypoglossal nucleus (tracheosyringeal, nXIIts) contains the motor neurons that innervate the syrinx in songbirds. It receives projections from telencephalic and midbrain nuclei that are necessary for song production. Its neurons concentrate androgens. The present study assesses the gross morphology of the hypoglossal nucleus in canaries. In this species song is more frequent, elaborate, and stereotyped in males than in females. Adult females respond to testosterone by developing a stereotyped song that is sung frequently. Song in male canaries is much more disrupted by damage on the left side of the song system than by damage on the right. We find anatomical correlates for each of these attributes in the nXIIts. This nucleus is 83% larger in males than in females. This is caused primarily by a sex difference in neuropil volume as there is no significant sex difference in the number of neurons in nXIIts. nXIIts grows by 34% in females given testosterone as adults. It is about 8% larger on the left than on the right in males, females, and females treated with testosterone. Sex differences are also found in the rostral (lingualis) portion of nXII, which controls muscles of the tongue, but there is no effect here of adult treatment with testosterone. Comparisons of these data with earlier measures of synaptic density and morphology in nXIIts suggest that the testosterone acts on this nucleus by inducing a modest increase in synapse numbers and by altering the efficacy of synapses in nXIIts. This contrasts with the effects of testosterone on n. robustus archistriatalis, a telencephalic component of the song system in which testosterone induces massive amounts of synaptogenesis.

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PubMed1991

Reassessing the mechanisms and origins of vocal learning in birds.

The most widely accepted hypothesis of vocal imitation in birds pre-dates many recent studies on the behavior, anatomy, physiology and cell biology of this phenomenon. It states that vocal learning involves two steps: (1) an auditory memory is laid down, and then (2) vocal output is modified until the auditory feedback it generates matches the model. This black-box model of vocal imitation disregards circuitry. We now know that the brain pathways for vocal learning in birds include a series of well-defined nuclei and projections. Some of these nuclei and projections develop late in ontogeny, at the time when auditory models are first acquired and imitated. We also know that the pathways involved in song production respond to sound, an observation that blurs the demarcation between what is an auditory and what is a motor circuit. These and other recent discoveries call for a reassessment of the mechanisms and origins of vocal learning in birds and mammals.

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PubMed1991

A comparative study of the behavioral deficits following lesions of various parts of the zebra finch song system: implications for vocal learning.

Song production in song birds is controlled by an efferent pathway. Appended to this pathway is a "recursive loop" that is necessary for song acquisition but not for the production of learned song. Since zebra finches learn their song by imitating external models, we speculated that the importance of the recursive loop for learning might derive from its processing of auditory feedback during song acquisition. This hypothesis was tested by comparing the effects on song in birds deafened early in life and birds with early lesions in either of two nuclei--Area X and the lateral magnocellular nucleus of the anterior neostriatum (LMAN). These nuclei are part of the recursive loop. The three treatments affected song development differently, as reflected by various parameters of the adult song of these birds. Whereas LMAN lesions resulted in songs with monotonous repetitions of a single note complex, songs of Area X-lesioned birds consisted of rambling series of unusually long and variable notes. Furthermore, whereas song of LMAN lesioned birds stabilized early, song stability as seen in intact birds was never achieved in Area X-lesioned birds. Early deafness also resulted in poorly structured and unstable song. We conclude that Area X and LMAN contribute differently to song acquisition: the song variability that is typical of vocal development persists following early deafness or lesions of Area X but ends abruptly following removal of LMAN. Apparently, LMAN plays a crucial role in fostering the kinds of circuit plasticity necessary for learning.

Aging↗

PubMed1991

Production and survival of projection neurons in a forebrain vocal center of adult male canaries.

Neurons are produced in the adult canary telencephalon. Many of these cells are incorporated into the high vocal center (nucleus HVC), which participates in the control of learned song. In the present work, 3H-thymidine and fluorogold were employed to follow the differentiation and survival of HVC neurons born in adulthood. We found that many HVC neurons born in September grow long axons to the robust nucleus of the archistriatum (nucleus RA) and thus become part of the efferent pathway for song control. Many of these new neurons have already established their connections with RA by 30 d after their birth. By 240 d, 75-80% of the September-born HVC neurons project to RA. Most of these new projection neurons survive at least 8 months. The longevity of HVC neurons born in September suggests that these cells remain part of the vocal control circuit long enough to participate in the yearly renewal of the song repertoire.

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PubMed1990

On variables that affect estimates of the true sizes and densities of radioactively labeled cell nuclei.

Tritiated thymidine has been widely used as a nuclear marker of cell birth. The true diameters and packing densities (nuclei/microns 3) of such radioactively labeled nuclei cannot be measured directly from tissue sections. Here we show that existing stereological corrections cannot be applied to data from radioactively labeled nuclei. We empirically measured the number of silver grains exposed by nuclei containing tritiated thymidine. The nuclei were separated from the photographic emulsion by known thicknesses of fixed, embedded avian telencephalon. The results of this experiment were used to develop an equation that estimates the number of silver grains exposed by a cell nucleus of any given diameter, containing a given amount of radioactive label, and located at any given distance from the photographic emulsion. The equation also allows one to calculate the probability that a label-containing nucleus will be correctly classified as labeled. Simulations of the equation revealed that not all label-containing nuclei are correctly classified by using commonly employed identification procedures and that larger nuclei are less likely to be correctly classified than smaller nuclei, given the same amount of label. The equation can be used to modify one class of existing stereological equations so as to be applicable to measurements of radioactively labeled nuclei. Finally, we discuss the assumptions and limitations of this modification.

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PubMed1990

Birth of projection neurons in adult avian brain may be related to perceptual or motor learning.

Projection neurons that form part of the motor pathway for song control continue to be produced and to replace older projection neurons in adult canaries and zebra finches. This is shown by combining [3H]thymidine, a cell birth marker, and fluorogold, a retrogradely transported tracer of neuronal connectivity. Species and seasonal comparisons suggest that this process is related to the acquisition of perceptual or motor memories. The ability of an adult brain to produce and replace projection neurons should influence our thinking on brain repair.

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PubMed1990

Song learning in birds: the relation between perception and production.

The vocal control system of oscine songbirds has some perplexing properties--e.g. laterality, adult neurogenesis, neuronal replacement--that are not predicted by common views of how vocal learning takes place. Similarly, we do not understand the relation between the direct pathway for the control of learned song and the recursive pathway necessary for song learning. Some of the paradoxes of the vocal system of birds may disappear once the relation between the perception and production of learned vocalizations is better understood. To some extent, perception and production may be two closely related states of a same system.

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PubMed1990

Proliferation "hot spots" in adult avian ventricular zone reveal radial cell division.

Neurogenesis in the adult avian brain is restricted to the telencephalon. New neurons originate in the ventricular zone (VZ) from cells that have not been identified. We mapped the position of [3H]thymidine-labeled cells in the walls of the ventricles of the adult canary brain. Labeled VZ cells were restricted to the telencephalon (lateral ventricles) and concentrated in "hot spots". The coincidence of these hot spots with regions rich in radial cells suggested that radial cells may be the cells undergoing mitosis. We used smears prepared from fragments of the VZ containing the hot spots to show directly that radial cells accumulate [3H]thymidine. In addition, grain counts at different survival times demonstrated that these cells divide. Hot spots of VZ cell division also coincided with sites of neuronal origin. We suggest that radial cell division may give rise to new neurons.

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