Volume Of Song Control Nuclei Research Articles

Testis-dependent and -independent effects of photoperiod on volumes of song control nuclei in American tree sparrows ( Spizella arborea)

Songbirds exhibit seasonal changes in the volumes of song control nuclei. Birds on long, spring-like days have larger nuclei than do birds on short, winter-like days. The mechanisms mediating volumetric changes have not been determined unequivocally, but testosterone (T) is probably involved. This study examined whether testicular factors are uniquely responsible for seasonal changes in the song system, or whether photoperiod has testis-independent effects. Male American tree sparrows were exposed to one of three photoperiodic conditions: (1) Photosensitive birds were retained on short days (8L:16D). Plasma T is rarely detected in such birds. (2) Photosensitive birds were moved from short days to long days (20L:4D) and photostimulated for three weeks. Photostimulation elevates circulating T in photosensitive birds. (3) Photorefractory birds were held at least four months on 20L:4D. Such birds seldom have detectable levels of T, even though they are on long days. In each condition, there were both intact and castrated birds. Castration typically removes circulating T in tree sparrows. The volumes of the high vocal center (HVC), nucleus robustus archistriatalis (RA), and area X were measured. Song nuclei were largest in intact photostimulated birds. Other long-day birds (i.e. castrated photostimulated, and intact and castrated photorefractory groups) had larger song nuclei than did short-day intact or castrated photosensitive birds and did not differ from each other. These data indicate that photoperiod has both testis-dependent and -independent effects on the volumes of song control nuclei.

Photoperiodic Condition Modulates the Effects of Testosterone on Song Control Nuclei Volumes in Male European Starlings

Songbirds possess a specialized network of brain nuclei mediating song learning and production, the song system. In many species, the nuclei in this circuit are larger in volume in males housed on long photoperiods than in males housed on short photoperiods. Exposure to long days initially leads to increases in circulating levels of testosterone (T) and it is generally believed that volume changes in the song system are controlled by T-dependent effects of photoperiod. However, a growing body of evidence indicates that photoperiod can also have T-independent effects on song system morphology. Previous work in our laboratory suggested that photoperiodic condition may modulate the effects of exogenous testosterone on the volumes of song control nuclei in male European starlings (Sturnus vulgaris). In the present report, we tested the hypothesis that photorefractoriness attenuates the effects of T on the volumes of song control nuclei in starlings. One group of long-day photorefractory males (16L:8D) was exposed to exogenous T and two groups of short-day photosensitive birds (8L:16D) were either implanted with T or blank (control) capsules. After 8 weeks, T levels were elevated in both groups of T-implanted birds and were undetectable in control animals. Volumes of the high vocal center were larger in T-implanted photosensitive birds than in both T-implanted photorefractory and control photosensitive males, which did not differ. These results suggest that photorefractoriness renders the song nuclei of starlings less sensitive to stimulatory effects of T. Plasticity in the starling song system appears to require the coordination of the appropriate hormonal milieu with a permissive photoperiodic condition.

Age‐ and behavior‐related variation in volumes of song control nuclei in male European starlings

There is considerable interindividual variation in the volumes of song control nuclei. Sex and physiological condition appear to contribute to these differences; however, these factors alone do not account for all of the variation. Studies have attempted to relate differences in song behavior (i.e., song repertoire size) to variation in song nucleus volume, but have met with mixed success. In this article, two studies are presented that used male European starlings (Sturnus vulgaris) to explore the relationship between song nuclei volumes and age-related differences in song behavior and interindividual variation in song behavior in adults. The results of the first study showed that song repertoire size and song bout length were significantly greater in older adult than in yearling males. In addition, the volumes of the high vocal center (HVC) and nucleus robustus archistriatalis (RA) were significantly larger in older adults than yearlings. Area X of the parolfactory lobe did not differ significantly in volume between the two age classes. In the second study, both HVC and RA volume correlated positively with song bout length but not repertoire size among adult birds. Based on these results a new hypothesis is presented that states that variation in song nuclei volumes in starlings relates more to the amount of song produced than to the number of song types stored in memory. © 1996 John Wiley & Sons, Inc.

Age- and behavior-related variation in volumes of song control nuclei in male European starlings.

There is considerable interindividual variation in the volumes of song control nuclei. Sex and physiological condition appear to contribute to these differences; however, these factors alone do not account for all of the variation. Studies have attempted to relate differences in song behavior (i.e., song repertoire size) to variation in song nucleus volume, but have met with mixed success. In this article, two studies are presented that used male European starlings (Sturnus vulgaris) to explore the relationship between song nuclei volumes and age-related differences in song behavior and interindividual variation in song behavior in adults. The results of the first study showed that song repertoire size and song bout length were significantly greater in older adult than in yearling males. In addition, the volumes of the high vocal center (HVC) and nucleus robustus archistriatalis (RA) were significantly larger in older adults than yearlings. Area X of the parolfactory lobe did not differ significantly in volume between the two age classes. In the second study, both HVC and RA volume correlated positively with song bout length but not repertoire size among adult birds. Based on these results a new hypothesis is presented that states that variation in song nuclei volumes in starlings relates more to the amount of song produced than to the number of song types stored in memory.

A direct comparison of the masculinizing effects of testosterone, androstenedione, estrogen, and progesterone on the development of the zebra finch song system

To assess which hormones are capable of masculinizing the neural song system of zebra finch hatchlings, we implanted female hatchlings with estrogen (estradiol [E2], 75 micrograms, n = 9), testosterone (T, 75-88 micrograms, n = 13), androstenedione (AE, 75 micrograms, n = 7), progesterone (P, 117 micrograms, n = 10), or nothing (Blanks, n = 10) and compared these to unimplanted males (n = 7). Implants, consisting of a hormone and Silastic mixture encased in polyethylene tubing, were placed under the skin of the breast on the day of hatching. Birds were killed when they were sub-adult (58 to 68 days old). We measured volumes of area X, the higher vocal center (HVC), and the robust nucleus of the archistriatum (RA); measured soma sizes in the lateral magnocellular nucleus of the neostriatum (lMAN), HVC, and RA; and counted RA neurons. E2 masculinized all measures in the song system and nearly sex-reversed the size of RA neurons. T masculinized volumes of nuclei and soma sizes but not the number or spacing of RA neurons. E2 was always at least as effective as T in masculinizing measures of the song system and was usually more effective. AE and P did not significantly masculinize any measure. These data suggest that E2 is more potent than aromatizable androgens or P in masculinizing the female song system in development and that the action of E2 alone may be sufficient to masculinize the volume of song control nuclei and the size and number of neurons.

Sexual dimorphism in the volume of song control nuclei in European starlings: Assessment by a Nissl stain and autoradiography for muscarinic cholinergic receptors

Previous studies have found that the volume of several song control nuclei is larger in male songbirds than in female songbirds. The degree of this volumetric sex difference within a given species appears to be systematically related to the degree of the behavioral sex difference. The largest volumetric differences have been reported in species in which the male sings and the female sings little, if at all, and the smallest sex differences in volume have been reported in species in which males and females both sing in nearly equal amounts. We compared the volume of three song control nuclei in male and female European starlings (Sturnus vulgaris), a species in which females are known to sing, though at a much lower rate than males. We investigated the volume of hyperstriatum ventrale, pars caudale, nucleus robustus archistriatalis, and area X of the lobus parolfactorius as defined with the use of a Nissl stain. In addition, we measured the volume of area X as defined by the density of muscarinic cholinergic receptors visualized by in vitro receptor autoradiographic methods. The volumes of all three of the song nuclei, as defined by Nissl staining, are significantly larger in males than in females. For area X, Nissl staining and receptor autoradiography indicate the same significant volumetric sex difference. The three nuclei are approximately one and one half to two times larger in males than in females, a degree of dimorphism that is intermediate to those reported for other species. Previous investigations of sex differences in the avian vocal control system have used only Nissl stains to define nuclear volumes. We demonstrate in this paper that receptor autoradiography can be used to assess dimorphisms in nuclear volume. Broad application of this approach to a number of neurotransmitter receptor systems will better characterize the dimorphisms in the song system, and therefore will provide greater insight into the neuroanatomical and neurochemical control of birdsong.

Changes in neuronal number, density and size account for increases in volume of song-control nuclei during song development in zebra finches

The caudal nucleus of the ventral hyperstriatum (HVc) and the robust nucleus of the archistriatum (RA) are two anatomically discrete brain regions that are known to be involved with song production in adult passerine birds. Both the HVc and RA increase greatly in volume during a restricted period of song development in male zebra finches, while brain regions not involved with song control show little or no increase in size. We report here that the increased volume of the HVc is attributable to an increase in the number of neurons during this period of song learning, whereas the growth of the RA is due to an increase in the somal size of neurons and a decrease in neuronal density. The pattern of results described is consistent with the idea that the HVc matures prior to the RA, and that the development of the RA may depend on the ingrowth of axons from the HVc and other song-control regions.