Endogenous oxytocin, cortisol, and testosterone in response to group singing.

Humans have sung together for thousands of years. Today, regular participation in group singing is associated with benefits across psychological and biological dimensions of human health. Here we examine the hypothesis that a portion of these benefits stem from changes in endocrine activity associated with affiliation and social bonding. Working with a young adult choir (n = 71), we measured changes salivary concentrations of oxytocin, cortisol, and testosterone from before and after four experimental conditions crossing two factors: vocal production mode (singing vs. speaking) and social context (together vs. alone). Salivary oxytocin and cortisol decreased from before to after the experimental manipulations. For oxytocin the magnitude of this decrease was significantly smaller after singing compared to speaking, resulting in concentrations that were significantly elevated after singing together compared to speaking together, after controlling for baseline differences. In contrast, the magnitude of the salivary cortisol decreases was the same across experimental manipulations, and although large, could not be separated from diurnal cycling. No significant effects were found in a low-powered exploratory evaluation of testosterone (tested only in males). At a psychological level, we found that singing stimulates greater positive shifts in self-perceived affect compared to speaking-particularly when performed together-and that singing together enhances feelings of social connection more than speaking together. Finally, measurements of heart rate made for a subset of participants provide preliminary evidence regarding physical exertion levels across conditions. These results are discussed in the context of a growing multidisciplinary literature on the endocrinological correlates of musical behavior. We conclude that singing together can have biological and psychological effects associated with affiliation and social bonding, and that these effects extend beyond comparable but non-musical group activities. However, we also note that these effects appear heavily influenced by broader contextual factors that shape social dynamics, such as stress levels, the intimacy of interactions, and the status of existing relationships.

Energetics of the acrobatic courtship in male golden-collared manakins (Manacus vitellinus).

In lek mating systems, females choose mates through indicators of quality, which males may exhibit by their performance of courtship displays. In temperate regions, displaying seasons are brief (one to two months), whereas in the tropics courtship seasons may be prolonged. Moreover, in temperate-breeding animals lekking behaviour can be energetically demanding, but little is known about the energy costs of lekking in tropical animals. Daily, over the course of a nearly seven-month-long breeding season, male golden-collared manakins (Manacus vitellinus) of Panamanian rainforests perform acrobatic courtship displays that markedly elevate heart rates, suggesting that they require high energy investment. Typically, animals of tropical lowland forests (such as manakins) exhibit a 'slow pace of life' metabolic strategy. We investigated whether male manakin courtship is indeed metabolically costly or whether the birds retain a low daily energy expenditure (DEE), as seen in other tropical species. To assess these questions, we calibrated manakin heart rate against metabolic rate, examined daily lek activity and, using telemetry, obtained heart rates of individual wild, lekking male manakins. Although metabolic rates peak during courtship displays, we found that males actually invest minimal time (only approx. 5 min d(-1)) performing displays. As a consequence, the DEE of approximately 39 kJ d(-1) for male manakins is comparable to other lowland tropical species. The short, intense bursts of courtship by these birds make up only approximately 1.2% of their total DEE. Presumably, this cost is negligible, enabling them to perform daily at their arenas for months on end.

Endocrine disrupters: a review of some sources, effects, and mechanisms of actions on behaviour and neuroendocrine systems.

Some environmental contaminants interact with hormones and may exert adverse consequences as a result of their actions as endocrine disrupting chemicals (EDCs). Exposure in people is typically a result of contamination of the food chain, inhalation of contaminated house dust or occupational exposure. EDCs include pesticides and herbicides (such as dichlorodiphenyl trichloroethane or its metabolites), methoxychlor, biocides, heat stabilisers and chemical catalysts (such as tributyltin), plastic contaminants (e.g. bisphenol A), pharmaceuticals (i.e. diethylstilbestrol; 17α-ethinylestradiol) or dietary components (such as phytoestrogens). The goal of this review is to address the sources, effects and actions of EDCs, with an emphasis on topics discussed at the International Congress on Steroids and the Nervous System. EDCs may alter reproductively-relevant or nonreproductive, sexually-dimorphic behaviours. In addition, EDCs may have significant effects on neurodevelopmental processes, influencing the morphology of sexually-dimorphic cerebral circuits. Exposure to EDCs is more dangerous if it occurs during specific 'critical periods' of life, such as intrauterine, perinatal, juvenile or puberty periods, when organisms are more sensitive to hormonal disruption, compared to other periods. However, exposure to EDCs in adulthood can also alter physiology. Several EDCs are xenoestrogens, which can alter serum lipid concentrations or metabolism enzymes that are necessary for converting cholesterol to steroid hormones. This can ultimately alter the production of oestradiol and/or other steroids. Finally, many EDCs may have actions via (or independent of) classic actions at cognate steroid receptors. EDCs may have effects through numerous other substrates, such as the aryl hydrocarbon receptor, the peroxisome proliferator-activated receptor and the retinoid X receptor, signal transduction pathways, calcium influx and/or neurotransmitter receptors. Thus, EDCs, from varied sources, may have organisational effects during development and/or activational effects in adulthood that influence sexually-dimorphic, reproductively-relevant processes or other functions, by mimicking, antagonising or altering steroidal actions.

Mating dominance amongst male Himalayan tahr: blonds do better.

In ungulates, rank order is determined by differences in weight, body size, weapon size and age. In the Caprini tribe (Bovidae: Caprinae), adult male Himalayan tahr are unique to show different coat colours, but no sexual dimorphism in weapons. A highly significant correlation between hair colour and rank order was found during the rut: males with a lighter coloured ruff dominated over darker ruffed ones, in both aggressive interactions and access to oestrus females. We studied colour-based dominance in relation to weight, age and testosterone levels, which establish the social rank in most ungulates. No differences in weight and testosterone concentrations were found between adult male colour classes, but males with paler ruffs were significantly younger than darker adult males. The distribution of physical traumas from fights confirmed that younger, lighter-coloured males had a higher rank than older, darker males, a pattern which is unusual amongst ungulates. Coat colour seems to work as a signal of rank in male-male aggressive interactions and it changes according to age, whereas the relevant physiological determinants deserve further research. Intrasexual male competition has not changed weapon size or shape in the Himalayan tahr, but ruff colours are apparently used to signal rank and dominance. Colour patterns of adult males may then be homologous to ritualised weapons, apparently being a unique feature of male tahr amongst mammals.

Hormonal influence on song structure and organization: the role of estrogen.

The development of song in songbirds is a complex phenomenon that involves memory and learning, sensorimotor integration, and neural and muscular maturation. Gonadal hormones are involved in each of these steps, as they influence the differentiation of the neural song system, the incorporation and survival of neurones, and the development of muscles used for song production. In young males the development of song, therefore, is closely linked to the secretion of testosterone by the testicles. Castration results in the development of incomplete or unstable songs, and hormone replacement leads to the development of crystallized or stable song. However, testosterone does not act solely as an androgen. The brain of songbirds contains high concentrations of the enzyme aromatase, which converts testosterone into estradiol. Estradiol then binds to estrogen receptors, which in the song system are found only in the nucleus HVC. This forebrain nucleus, also called the "master nucleus," codes for the syntactic structure of song, i.e. for the particular combination of simple elements-syllables-that characterize the song of an individual. In this paper, we will review our studies on the role of estrogen in guiding the organization of song in canaries.

Difficulties and special issues associated with field research in behavioral neuroendocrinology.

Classical behavioral neuroendocrinology has focused on a limited number of domestic mammals and birds. The model systems used in these studies represent a very small proportion of the diversity of hormone-behavior interactions found in nature. In the last three decades, an increasing number of researchers have concentrated their efforts on studying behavioral neuroendocrinology of wild animals. Field behavioral neuroendocrinology presents a series of challenges ranging from the design of the experiments to sample preservation and transportation. The constraints of field conditions limit the number of factors that can be controlled for and the questions that can be addressed. On the other side, many behaviors can be studied only in the field, and only a few species can be kept in captivity. Thus, field studies are necessary to understand the complexity and variety of interactions between hormones, brain, and behavior. In this article, we will review some of the peculiarities and challenges of field behavioral neuroendocrinology, including solutions for some of the most commonly encountered technical issues.

Testosterone regulates the activity and expression of aromatase in the canary neostriatum.

The estrogen synthesizing enzyme, P450 aromatase, plays a critical role in the regulation of vertebrate sexual behavior. Songbirds differ from other avian species in the distribution and expression of aromatase in the telencephalon. The highest concentration of aromatase in the songbird brain is found in the caudomedial neostriatum (NCM). This area surrounds the only nucleus of the neural song system that contains estrogen receptors, the high vocal center (HVC). It has been suggested that estrogen produced in NCM via aromatization of circulating testosterone (T) is involved in song development and adult song plasticity. The modalities of regulation of aromatase in NCM are not well understood, and some studies suggest that in NCM, unlike in the preoptic-hypothalamic areas, aromatase is not regulated by androgen and/or estrogen. In this work, we studied whether the treatment of female canaries with T, which induces the development of malelike song and the masculinization of the song system, also induces an increase in the expression and activity of aromatase in NCM. Our results show that both the expression and activity of aromatase in NCM increase in female canaries following T treatment. This study provides the first direct evidence that T regulates telencephalic aromatase in songbirds, and suggests that an increase in estrogen production in NCM might be functional in neural and behavioral plasticity during phases of song organization.

Aromatase inhibition reduces specifically one display of the ring dove courtship behavior.

The courtship behavior of the male ring dove (Streptopelia risoria) combines aggressive displays (chasing, bowing) and nest-oriented displays (nest soliciting). Aggressive displays depend on circulating testosterone, whereas nest soliciting is estrogen-dependent and appears to depend on the aromatization of androgen into estrogen within the brain. The present work tested the hypothesis that aromatase specifically modulates the nest soliciting display in intact male ring doves. Males were tested for courtship behavior with receptive females before and after being implanted with micro-osmotic pumps containing Fadrozole, a nonsteroidal aromatase inhibitor, or saline. Fadrozole at the higher dose reduced estrogen-dependent nest soliciting but did not affect androgen-dependent chasing and bowing. These results support the hypothesis that aromatase modulates nest soliciting in male ring doves, and provide further evidence for separate hormonal control of different courtship displays in this species.

Seasonal expression of androgen receptors, estrogen receptors, and aromatase in the canary brain in relation to circulating androgens and estrogens.

Songbirds have a complex neural network for learning and production of song, namely the neural song system. Several nuclei of the song system contain androgen receptors (AR), and the neostriatal nucleus HVc also contains alpha type estrogen receptors (ER). Many songbird species show seasonal changes in both song and the neural song system that are correlated with seasonal variations in the circulating levels of gonadal steroids. However, there is increasing evidence that the sensitivity of the song system to gonadal steroids also changes seasonally. This could involve changes in the expression and activity of steroid receptors and steroid-metabolizing enzymes, such as the estrogen-synthesizing enzyme aromatase (AROM). The seasonal regulation of brain AR, ER, and AROM has not been studied before in the same individual songbirds. In this work, we compared plasma levels of androgens and estrogens, the expression level of AR-, ER-, and AROM-mRNA in the telencephalon, and brain AROM activity in male canaries between autumn (November) and spring (April) periods of high singing activity. Plasma levels of androgens and estrogens were higher in April than in November. The expression level of ER in HVc was higher in November than in April. In contrast, the expression level of AROM in the caudomedial neostriatum was higher in April than in November. However, we found no seasonal differences in the level of expression of AR and the volume of HVc as delimited by AR expression. Thus, AR expression in HVc was not correlated with circulating androgen levels. This study shows that both steroid-dependent and -independent seasonal factors regulate the action of gonadal hormones on the song system. In addition, we report a new site of AROM expression in the songbird brain, the nucleus interfacialis.

Distribution of aromatase, estrogen receptor, and androgen receptor mRNA in the forebrain of songbirds and nonsongbirds.

Androgens and estrogens are crucial for the differentiation and function of the vocal control system of songbirds. A major source of estrogens in songbirds is the cerebral aromatization of circulating testosterone by aromatase (ARO). In the vocal control system, songbirds have a unique estrogen receptor (ER)-containing area, the nucleus hyperstriatalis ventrale pars caudale (HVC) of the caudal neostriatum. Work in the zebra finch has demonstrated ARO expression adjacent to but not in the HVC. Compared with other songbirds, such as the canary, the HVC of adult zebra finches contains only few ERs. To determine whether the disjunctive distribution of ERs and ARO in the forebrain is a songbird-specific feature, the authors investigated ARO and ER mRNA expression in songbirds (canary, house sparrow, and zebra finch) and in nonsongbirds (budgerigar, ring dove, swift, grey partridge, and barn owl) of five avian orders. In addition, the coexpression of androgen receptor (AR) and ARO mRNAs was studied. Preoptic hypothalamic areas showed similar expression of ARO in all species. In the caudal neostriatum, ARO, AR, and ER transcripts were found only in songbirds. ARO and ER mRNA expression in the caudal forebrain was spatially separated, i.e., the HVC contained ER mRNA but very little or no ARO mRNA, and the caudomedial neostriatum contained high levels of ARO mRNA but few if any ERs. ARO and AR mRNAs, however, were coexpressed in the caudomedial neostriatum. The coexpression of ARO mRNA with AR mRNA but not with ER mRNA was found in further brain areas, such as the nucleus posterior lateralis hypothalami. The area-specific coexpression of AR, ER, and ARO suggests various possibilities for the steroid-dependent regulation of ARO and for the role of ARO in controlling AR- and ER-dependent mechanisms.

Sexually selected vigilance behaviour of the grey partridge is affected by plasma androgen levels

In the grey partridge, Perdix perdixvigilance and calling activity are two sex-dimorphic behaviours that are critical for mate choice. To ascertain whether circulating levels of testosterone directly affect vigilance (i.e. the occurrence of upright alert posture), we compared vigilance scores of testosterone-implanted versus control males both during the normal activity of the flock and after the passage of a raptor silhouette; in the latter case, the calling activity was also recorded. Hormone-treated males were more vigilant than controls in both experimental situations. Vigilance was correlated with calling rate. Testosterone seems to act as a link, relating conspicuous behaviours involved in sexual selection to male quality and physical condition, because of the costs of having high levels of both signalling and androgens.Copyright 1997 The Association for the Study of Animal Behaviour1997The Association for the Study of Animal Behaviour

Regional myocardial blood flow in stable angina pectoris associated with isolated significant narrowing of either the left anterior descending or left circumflex coronary artery.

Myocardial perfusion measurements were obtained with positron emission tomography under basal conditions and after intravenous dipyridamole infusion (0.56 mg/kg over 4 minutes) to determine if myocardial perfusion is maximized in areas of resting wall motion abnormalities in patients with stable angina. Thirty-three patients with no history of myocardial infarction, and with coronary stenosis > 50% involving the left anterior descending (n = 24) and left circumflex (n = 9) coronary arteries were evaluated. Quantitative perfusion images were recorded twice in each subject using nitrogen-13 ammonia at baseline and after intravenous administration of dipyridamole. Computer-assisted analysis of left ventriculograms showed abnormal wall motion in the stenosis-related regions in 16 patients (group 1), and normal regional function in 17 (group 2). The flow values in the anterior and posterolateral wall were considered to reflect left anterior and left circumflex coronary artery flow, respectively. Quantitative angiography showed that coronary stenosis severity was higher in group 1 than in group 2 (cross-sectional area reduction 94 +/- 7% vs 87 +/- 11%; p < 0.05). Resting blood flow in the stenosis-related areas was significantly lower than in contralateral regions in group 1 (0.66 +/- 0.19 vs 0.77 +/- 0.26 ml/min/g; p < 0.05), but not in group 2 (0.73 +/- 0.18 vs 0.78 +/- 0.21 ml/min/g; p = NS). Dipyridamole significantly (p < 0.01) increased myocardial blood flow in both stenotic and remote regions in both groups 1 (0.95 +/- 0.41 vs 1.57 +/- 0.70 ml/min/g) and 2 (1.54 +/- 0.53 vs 2.01 +/- 0.84 ml/min/g).(ABSTRACT TRUNCATED AT 250 WORDS)

Simultaneous in vitro and in vivo validation of nitrogen-13-ammonia for the assessment of regional myocardial blood flow.

Measurement of myocardial blood flow by 13NH3 relies heavily on the assessment of both the input function and the variable tissue extraction fraction. In six open-chest dogs, myocardial and arterial 13NH3 activity was measured both by in vitro sampling and by in vivo positron emission tomography (PET). Regional myocardial blood flow was forced to vary in the range 0.2-5 ml/min/g and actual values were assessed by in vitro counting of 153Gd microspheres. The ammonia input function was processed by: (a) total curve integration; (b) curve integration for 2 min; (c) integral of a fitted curve (gamma variate in vivo and exponential of the downslope in vitro). Method C brought to regional flow values which best approximated microspheres data. The in vitro correlation allows for correcting in vivo values for the flow-dependent extraction fraction. The method can be easily applied for regional myocardial blood flow measurements with PET in human studies.

High-performance liquid chromatographic separation of iodoamino acids for tracer turnover studies of thyroid hormones in vivo.

A reversed-phase high-performance liquid chromatographic technique was developed to separate radioiodinated thyroxine (T4), 3,5,3'-triiodothyronine (T3), 3,3',5'-triiodothyronine (rT3) and two diiodothyronines (3,3'-T2 and 3',5'-T2), in extracts from either serum or urine. Chromatography was performed with 10-micron C18 silica gel, packed in a glass column (3 X 300 mm); the mobile phase was methanol-water (55:45) adjusted to pH 3 with H3PO4, at a flow-rate of 1.2 ml/min and a pressure of 2800 p.s.i. The results demonstrate the ability of the system to yield a clear-cut separation of the iodothyronines involved in in vivo turnover studies, i.e., T4, T3, rT3, and the two T2 compounds together.

Coagulation factor XIII in patients with acute and chronic renal disease.

The behavior of coagulation factor XIII (fibrin-stabilizing factor, FSF) was studied in patients with renal disease. Specific antiserum against the active subunit (FSFA) was employed to set up a method for the direct measurement of the active fraction in plasma, according to the electroimmunodiffusion technique. The plasma FSFA levels were measured in the following patients: (I) 31 patients with chronic renal disease and serum creatinine not higher than 1.5 mg/dl; (II) 41 patients with chronic renal failure on conservative therapy; (III) 53 uremic patients on maintenance hemodialysis; (IV) 10 patients with acute renal failure. FSFA concentration (93.3 +/- 17.6% of a reference plasma in a group of 15 healthy controls) was found to be significantly higher than normal in the patients with chronic renal disease and serum creatinine lower than 1.5 mg/dl (127 +/- 39.8%, p less than 0.005). The FSFA levels were similarly increased in the 41 patients with chronic renal failure on conservative management (134.9 +/- 35.8%, p less than 0.001), and in the 53 end-stage uremics on maintenance hemodialysis (132.8 +/- 29.5%, p less than 0.001). Whereas, FSFA concentration was found to be markedly reduced in the 10 patients with acute renal failure (35.8 +/- 14.6%, p less than 0.001). In the patients with chronic renal disease (groups I, II, and III) plasmatic FSFA higher in those patients with serum triglycerides above the upper normal limit, and a significant positive correlation was found between serum triglycerides and FSFA plasma levels.