Communication goes multimodal.
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Biomedical subjects
Publications and source records attributed to P Marler.
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Song learning in white-crowned sparrows, Zonotrichia leucophrys, involves three steps: memorization of external models, song practice and selection of a song from the practiced repertoire for crystallization. These three events occur in a sequential and predictable order during the first year of life in captive sparrows. To study the external regulation of these events, we raised nestling sparrows under conditions in which photoperiod and tutor exposure were manipulated. We measured plasma testosterone concentration twice a month to study its role in the mediation of vocal learning. We conclude that the timing of song memorization is relatively impervious to photoperiodic manipulation. Song practice and crystallization, however, were readily influenced by both photoperiod and tutor exposure. We suggest that low testosterone concentrations permit acquisition at an older age than would normally occur, and confirm that testosterone propels the transition to production of crystallized song, but is not required for the onset of song practice. Copyright 1998 The Association for the Study of Animal Behaviour.
The anterior forebrain pathway of the avian song system is involved in juvenile song learning, but its function in adult song behavior is not known. This report uses lesions to study the role of a particular forebrain nucleus, IMAN, in the seasonal regeneration of song in adult white-crowned sparrows (Zonotrichia leucophrys oriantha). White-crowned sparrows, even when acoustically isolated as juveniles, crystallize a single song which they maintain throughout adulthood. The lateral portion of the magnocellular nucleus of the anterior neostriatum (1MAN) was lesioned bilaterally in adult males maintained on short days (8 h of light). Daylength was increased to 16 h following the surgeries, and all birds were recorded in the post-lesion singing season. Lesioned birds showed a large decrease in song note frequency following the lesions, significantly larger than did intact, age-matched controls. Further changes were seen in the post-lesion songs of seven of 11 successfully lesioned males. These changes included variability in song pattern, loss of frequency control and addition of new notes, some of which had been practiced during juvenile song development. These changes seemed especially large in birds that had either been acoustically isolated or had not fully copied a tape-tutor song during juvenile song development. These results are the first to indicate that the motor memories for song elements that had been practiced and discarded early in life are retained, and they suggest that 1MAN affects seasonal song expression by selectively reinforcing a particular song pattern.
In behavior reminiscent of the responsiveness of human infants to speech, young songbirds innately recognize and prefer to learn the songs of their own species. The acoustic and physiological bases for innate recognition were investigated in fledgling white-crowned sparrows lacking song experience. A behavioral test revealed that the complete conspecific song was not essential for innate recognition: songs composed of single white-crowned sparrow phrases and songs played in reverse elicited vocal responses as strongly as did normal song. In all cases, these responses surpassed those to other species' songs. Although auditory neurons in the song nucleus HVc and the underlying neostriatum of fledglings did not prefer conspecific song over foreign song, some neurons responded strongly to particular phrase types characteristic of white-crowned sparrows and, thus, could contribute to innate song recognition.
We describe a new assay for measuring the acquisition phase of song learning in birds and compare it with the standard method of comparing a pupil's imitation to his tutor's song. Juvenile male and female white-crowned sparrows, Zonotrichia leucophrysgive call notes in response to playback of species-typical song. After 10-day periods of tape tutoring conducted in the first 50 days of life, subjects gave significantly more calls in response to tutor songs than to novel songs. An identical procedure conducted at a later age yielded a different result. Birds did not distinguish tutor from novel songs in winter, after the sensitive phase for song acquisition. Males gave significantly more calls to those tutor songs that they subsequently sang the next spring than to tutor songs they did not reproduce. Assuming that the call assay correlates with long-term storage of songs in females as it does in males, these results also indicate that the sensitive phase for song acquisition is shorter in females than in males of this species.1997The Association for the Study of Animal Behaviour
In an effort to prepare 3,4-methylene-dioxyphenyl-(S)-isopropanol from 3,4-methylene-dioxyphenylacetone, an initial screen of microbes indicated that Candida famata could catalyze this reaction efficiently at low substrate concentration. A dilute, large-scale process was developed to provide experimental material for the chemical synthesis to be explored. However, the productivity number of this process [0.134 g product (g wet weight cells)-1 day-1 was too low to be practical. C. famata was also extremely sensitive to concentrations of both the ketone and the alcohol greater than 2 g/l. A more extensive screen of yeast and fungi revealed that Zygosaccharomyces rouxii was more tolerant to higher substrate concentrations and had a higher productivity number [0.8 g (g wet weight cells)-1 day-1]. These characteristics suggested that Z. rouxii could be used in a large-scale process at high substrate concentrations.
Research on avian song learning has traditionally been based on an instructional model, as exemplified by the sensorimotor model of song development. Several large-scale, species-wide field studies of learned birdsongs have revealed that variation is narrowly restricted to certain aspects of song structure. Other aspects are sufficiently stereotyped and so widely shared by species' members that they qualify as species-specific universals. The limitations on natural song variation are difficult to reconcile with a fully open, instructive model of song learning. An alternative model based on memorization by selection postulates a system of innate neural templates that facilitate the recognition and rapid memorization of conspecific song patterns. Behavioral evidence compatible with this model includes learning preferences, rapid conspecific song learning, and widespread ocurrence of species-specific song universals that are recognized innately but fail to develop in songs of social isolates. A third model combines instruction, in the memorization phase, with selection during song production. An overproduced repertoire of plastic songs previously memorized by instruction is winnowed by selection imposed during social interactions at the time of adult song crystallization. Selection during production is well established as a factor in the song development of several species, in the form of action-based learning. The possible role of selective processes in song memorization merits further neurobiological investigation.
Bird song is a complex, learned behavior. Vocal learning in sparrows involves several different processes that occur in a distinct temporal pattern over the course of the first year of life. Songs are acquired without practice during a sensitive period within the first 3 months of life and rehearsal of the acquired song does not begin until 7 or 8 months of age. The function of the storage period between song acquisition and production is not known. We set out to investigate its significance by administering testosterone, known to stimulate production of adult song, to birds at 100 days of age after song acquisition was completed but some 5 months prior to normal song onset. Most testosterone-treated birds produced abnormal songs resembling those of males raised in acoustic isolation suggesting that, in sparrows, events occurring during the storage phase play a significant role in vocal learning.
Bird song is a model system for study of the neurobiology, development, and functions of learned vocal communication signals. Research on avian song learning has been dominated by an instructive model of learning--the sensorimotor model. Developmental plasticity is assumed to be based on the incorporation of novel material into the song repertoire. Experimental evidence now indicates an alternative form of plasticity operating in harmony with sensorimotor learning, based on principles of selection, rather than instruction. Song dialects, a common consequence of vocal learning, can be achieved by overproduction of previously memorized songs and selective attrition of those that fail to match the dialect of interacting males. These distinct processes of learning have important implications for study of the neural substrates underlying song production and perception and for efforts to interpret patterns of geographic variation in song.
In previous studies, androstenedione (AE) replacement therapy restored the highest levels and intensities of courtship song displays in castrated male zebra finches of any hormone tested. Furthermore, female zebra finches responded strongly to AE-treated males and preferred intact males given small AE implants to unsupplemented males. In this study, we asked whether AE treatment might alter song structure, since male song is an important cue in mate choice by female zebra finches. Songs of adult males were recorded. The males were then castrated and given AE therapy and recorded again. No differences were found between the courtship or undirected songs males sang before castration and after AE treatment. As in previous studies, the structure of a male's courtship song differed significantly from his undirected song, and the structural differences between these two song types were not altered by AE treatment.
Behavioral studies of song learning in birds continue to raise new problems for neuroethological investigation. Evidence is emerging for a new form of vocal plasticity, called 'action-based learning'. Motor patterns are overproduced during a particular phase of ontogeny, and are then subjected to attrition and selective reinforcement by various kinds of social stimulation as the young bird matures. This form of learning, analogous to operant conditioning, can occur at phases of development when the more traditional form of 'memory-based learning' is no longer possible. There is evidence that different physiological mechanisms are involved in the development and the maintenance of mature singing, with a transition occurring at the time of song crystallization. Neural events associated with the developmental stabilization of motor patterns are worthy of more study.
The high degree of developmental plasticity displayed by the songs of oscine birds makes them appropriate subjects for research on the etiology and neurobiology of vocal learning. Strong individual differences and learned local dialects are common. The readiness to acquire new songs appears to persist throughout life in some species and is restricted to relatively short sensitive periods in others. Learning can occur with remarkably few exposures to song. Mimicry of other species occurs but, given a choice, there is a tendency to favour conspecific songs. Evidence is presented for two kinds of vocal learning, one 'memory-based', the other 'action-based.' Subsong and 'plastic song' phases of motor development appear to be obligatory steps in the ontogeny of learned songs. A case is made that acquisition and production should be viewed as distinct phenomena with different physiological correlates. Research on behavioural development is closely associated with studies of the physiology of development. The two are mutually synergistic, and the synergism is well displayed in research on song learning in birds. This review of some of the characteristics of avian vocal learning as derived from behavioural studies, indicates lacunae in our knowledge about the ethology of song learning, and suggests how the comparative study of vocal development can pave the way for new insights into the underlying neurobiology.
Research on the ways in which different species of birds learn to sing is used to illustrate the necessity of taking innate factors into account in studies of behavioral development. Experiments on two species of songbirds are described that reveal innate species differences in responsiveness to tape-recorded songs. Conspecific songs are favored over those of other species. These patterns of innately varying responsiveness provide a basis for the development, not of stereotyped behavior, but of variable, individually learned behavior. The viewpoint is presented that mechanisms that differ innately from species to species, some with general functions, others specialized for particular ontogenetic assignments, provide the necessary substrates with which experience interacts.
We tested the ability of 1-year-old European starlings (Sturnus vulgaris) to acquire songs while in different physiological states. Photorefractory males, with low testosterone levels, learned songs as completely and as accurately as photosensitive males in full reproductive condition. This indicates that song acquisition in 1-year-old males does not depend on high levels of androgens. The ability to learn songs during the nonbreeding season may reflect the high song output of potential tutors through most of the year, including the photorefractory period, and may facilitate increases in song repertoire size in adulthood.
A fundamental issue in perception and communication is how continuously varying stimuli are partitioned into discrete categories. In swamp sparrow songs, note duration is a critical feature distinguishing two note categories with different roles in song construction. Pairs of songs with initial notes from different categories contrast more in their effects on territorial males than song pairs with initial notes differing by the same amount but taken from within one note category. The results indicate categorical perception by wild swamp sparrows.
During song learning in birds, neurons are added to some song nuclei and lost from others. Previous studies have been unable to distinguish whether these neural changes are uniquely associated with memorizing a song model (sensory acquisition) or vocal practice (sensorimotor learning). In this study we measured changes in neuron number within song nuclei of swamp sparrows, a species in which the two phases of song learning are nonoverlapping. Male swamp sparrows were collected as hatchlings and tape-tutored from approximately 22 to 62 days of age. Swamp sparrows memorize about 60% of their song material during this period, but do not begin practicing this learned material until approximately 275 days of age. Birds were sacrificed at 23, 41, 61, 71, 274, or 340 days of age. During sensory acquisition, neuron number increased drastically in both the caudal nucleus of the ventral hyperstriatum (HVc) and Area X. The period of sensorimotor learning was not associated with any further changes in neuron number within these regions. We were unable to detect any significant changes in neuron number within the magnocellular nucleus of the neostriatum or the robust nucleus of the archistriatum during either stage of song learning. These results raise the possibility that ongoing addition of HVc and Area X neurons may encourage, and thereby temporally restrict, song acquisition.
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