Ring Doves Research Articles

Syringeal muscles fit the trill in ring doves (Streptopelia risoriaL.)

In contrast to human phonation, the virtuoso vocalizations of most birds are modulated at the level of the sound generator, the syrinx. We address the hypothesis that syringeal muscles are physiologically capable of controlling the sound-generating syringeal membranes in the ring dove (Streptopelia risoria) syrinx. We establish the role of the tracheolateralis muscle and propose a new function for the sternotrachealis muscle. The tracheolateralis and sternotrachealis muscles have an antagonistic mechanical effect on the syringeal aperture. Here, we show that both syringeal muscles can dynamically control the full syringeal aperture. The tracheolateralis muscle is thought to directly alter position and tension of the vibrating syringeal membranes that determine the gating and the frequency of sound elements. Our measurements of the muscle's contractile properties, combined with existing electromyographic and endoscopic evidence, establish its modulating role during the dove's trill. The muscle delivers the highest power output at cycle frequencies that closely match the repetition rates of the fastest sound elements in the coo. We show that the two syringeal muscles share nearly identical contraction characteristics, and that sternotrachealis activity does not clearly modulate during the rapid trill. We propose that the sternotrachealis muscle acts as a damper that stabilizes longitudinal movements of the sound-generating system induced by tracheolateralis muscle contraction. The extreme performance of both syringeal muscles implies that they play an important role in fine-tuning membrane position and tension, which determines the quality of the sound for a conspecific mate.

Comparative Study of the Activity/Rest Rhythms in Young and Old Ringdove (Streptopelia Risoria): Correlation with Serum Levels of Melatonin and Serotonin

Aging is characterized by changes in the circadian rhythms of melatonin, serotonin, and sleep/wakefulness, alterations that affect sleep quality. The authors studied the circadian rhythms of serotonin and melatonin in young and old ringdoves (Streptopelia risoria) (2–3 and 10–12 yrs old, respectively), animals that are characterized by being monophasic and active by day, like humans. The aim was to correlate the indole rhythms with the animals' activity/rest periods. The animals were kept under a 12∶12 h light/dark cycle, fed ad libitum, and housed in separate cages equipped for activity recording. Activity pulses were recorded with one actometer per animal (two perpendicular infrared transmitters) and were logged every 15 min by a computer program (DAS 16) throughout the experiment. Melatonin was measured by radioimmunoassay and serotonin by ELISA at intervals of 3 h (from 09∶00 to 18∶00 h) and 1 h (from 21∶00 to 06∶00 h), respectively. The results showed a reduction in nocturnal vs. diurnal activity of 89% and 61% in the young and old animals, respectively, with 100% considered to be the diurnal activity of each group. The amplitude of a cosine function fit to the melatonin concentrations of the old animals was half that of the young birds. The acrophase and nadir were at 02∶00 and 14∶00 h in the young and 01∶00 and 13∶00 h in the old animals, respectively. The amplitude of the corresponding cosine function fit to the serotonin concentrations in the old birds was one‐third that of the young animals. The acrophase and nadir were at 15∶00 and 03∶00 h in the young and 16∶00 and 04∶00 h in the old animals, respectively. For both melatonin and serotonin, the concentrations in the young animals were significantly higher than in the old at most of the measurement times. There was a clear negative correlation between the circadian rhythms of activity and the serum melatonin levels in both young and old animals. The equivalent correlation for serotonin was positive, and stronger in the case of the young animals. The results suggest a possible relationship between the observed decline in the amplitude of the old animals' melatonin and serotonin rhythms and the lower percentage reduction in their nocturnal relative to diurnal activity pulses compared to the young animals. In conclusion, the circadian rhythms of melatonin and serotonin undergo alterations with age that could be involved in the changes in age‐associated sleep.

Patterns of Fos-like immunoreactivity in the brains of parent ring doves (Streptopelia risoria) given tactile and nontactile exposure to their young.

Neuronal activation was examined by fos immunohistochemistry in ring doves (Streptopelia risoria) reunited with their young after overnight separation. In an initial study, squab-exposed parents showed more fos immunoreactivity (ir) in the preoptic area (POA) and lateral hypothalamus (LH) than squab-deprived parents. In a 2nd study, parents allowed free access to young and those separated from young by a wire mesh partition showed more fos-ir in the POA, LH, and lateral septum than box-exposed controls. Contact with young also increased fos-ir in the medial preoptic nucleus and bed nucleus of the stria terminalis, but noncontact exposure did not. Conversely, nontactile squab exposure stimulated more fos-ir in the POA than did free access to young, which suggests POA involvement in appetitive aspects of parenting.

Age-related changes in circadian rhythm of serotonin synthesis in ring doves: Effects of increased tryptophan ingestion

Alterations in the function of the hypothalamic suprachiasmatic nucleus (SCN) with age have been reported. As serotonin is an important regulator of the circadian clock located in SCN, this work studied the changes produced in the synthesis of serotonin with age using the accumulation of 5-HTP after decarboxylase inhibition as a measure of serotonin synthesis in the brain in vivo, in young and old ring doves at the onset of lights-on and lights-off. A diurnal cycle in tryptophan hydroxylation was observed in young animals, with an increased daylight synthesis and metabolism of 5-HT in hippocampus, neostriatum and hypothalamus. A single dose of melatonin (1 mg/kg, i.p., 1 h) at lighttime produced an inhibitory effect on the synthesis of 5-HT. In contrast, differences in 5-HT synthesis and metabolism between day and night dissappeared in old animals indicating an absence of a circadian rhythm in 5-HT synthesis and metabolism. The administration of l-tryptophan (240 mg/kg, i.p.) strongly increased the 5-HT synthesis in young animals only during lights-off time while it increased in old ones irrespective of the administration time. These results suggest that the supplemental administration of tryptophan might aid to improve the descent in 5-HT that normally occurs, as animals get old.

Aromatase inhibition abolishes courtship behaviours in the ring dove (Streptopelia risoria) and reduces androgen and progesterone receptors in the hypothalamus and anterior pituitary gland

The aim of this study was to determine in the ring dove, the effects of aromatase inhibition on the expression of aggressive courtship and nest-soliciting behaviours in relation to the distribution of cells containing immunoreactive androgen (AR) and progesterone (PR) receptor in the hypothalamus and pituitary gland. Isolated sexually experienced ring doves were transferred in opposite sex pairs to individual breeding cages, and then injected with the aromatase inhibitor, fadrozole (four males and four females), or saline vehicle (four males and four females) for 3 days at 12 hourly intervals. Saline-injected control males displayed aggressive courtship behaviours (bow-cooing and hop-charging) and nest-soliciting throughout the study, and control females displayed nest-soliciting. By day 3, fadrozole treatment resulted in the disappearance of all these behaviours and in a decrease or disappearance of AR and PR in the anterior pituitary gland, and in the nucleus preopticus paraventricularis magnocellularis (PPM), nucleus preopticus medialis (POM), nucleus hypothalami lateralis posterioris (PLH), and ventral, lateral and dorsal nucleus tuberalis in the hypothalamus (VTu, LTu, DTu). In the nucleus preopticus anterior (POA), fadrozole treatment decreased AR in both sexes and decreased PR in females but not in males. Cells containing co-localized nuclear AR and PR were found in all hypothalamic areas examined, and in the anterior pituitary gland. Fadrozole is suggested to reduce the local availability of estrogen required indirectly for the induction of AR, and except in cells containing PR in the male POA, for the direct induction of PR. It is suggested that aggressive courtship behaviour is terminated by "cross talk" between aromatase-independent PR and aromatase-dependent AR co-localized in neurons in the POA. Aromatase-independent PR may increase in the male POA in response to visual cues provided by a partner. Aromatase-dependent PR in the POM, and basal hypothalamus may play a role in the facilitatory effect of progesterone on estrogen-induced nest-orientated behaviours.

Melatonin, lipid peroxidation, and age in heterophils from the ring dove (Streptopelia risoria)

Numerous recent studies have shown the ability of physiological as well as all pharmacological concentrations of melatonin to prevent oxidative stress. We have found that incubating avian heterophils from young birds with a pharmacological concentration of 100 μM (23 × 106 pg/ml) melatonin reduced superoxide anion levels by modulating the activity of superoxide dismutase while also enhancing phagocytosis. There was also a decline in lipid peroxidation levels with both physiological and pharmacological concentrations of this indolamine.In the present work, we evaluated malonaldehyde (MDA) levels as an indicator of lipid peroxidation (both basal and antigen-induced) in young and old animals (ring doves) at different times of day (16:00 and 00:00) and with two incubation times (15 and 60 min). The lipid peroxidation was also measured in heterophils from old animals, incubated with the physiological concentrations of melatonin measured in young animals (50 and 300 pg/ml, diurnal and nocturnal, respectively). The results, expressed as nmol MDA/mg protein, show that MDA levels were higher in heterophils of old animals than in the young birds in all the experimental groups studied at both 16:00 and 00:00 (00:00 is the time at which the lowest peroxidation levels were obtained). Incubation with melatonin was found to reduce MDA levels, with the maximum reduction being after the 60 min incubation time and the nocturnal melatonin concentration. At both concentrations (diurnal and nocturnal), melatonin also counteracted the enhancement of MDA levels caused by latex beads, with the effect being greater at the longer incubation time. In conclusion, the results are further evidence of the antioxidant effect of melatonin even at physiological concentrations, and suggest its utility as a therapeutic agent in some pathological processes associated with age.

A New Model for Sexual Conditioning: The Ring Dove (Streptopelia risoria).

Sexual conditioning investigations have primarily used rats and domestic quail as subjects. Although much has been learned from such experiments, the relative simplicity of rat and quail reproductive behavior prohibits investigation of certain experimental questions about sexual conditioning. In contrast, the reproductive behavior of male and female ring doves (Streptopelia risoria) is complex, involving courtship, bonding, and parental care. In the present experiment, male ring doves that were presented with a visual conditioned stimulus paired with access to their pair-bonded mate (the unconditioned stimulus) showed sexual conditioned responding in the presence of the conditioned stimulus. These results represent the first evidence of sexual conditioning in ring doves and illustrate the potential of ring doves as a useful model for future sexual conditioning investigations.

Neuronal activation related to auditory perception in the brain of a non-songbird, the ring dove

In songbirds, parrots, and hummingbirds, which need to learn their songs, exposure to conspecific song leads to increased expression of the immediate early gene (IEG) ZENK in a number of forebrain regions, including the caudomedial nidopallium (NCM) and the caudomedial mesopallium (CMM). Here we investigated the pattern of IEG expression in response to auditory stimulation in the brain of the ring dove (Streptopelia risoria), a non-songbird that does not need to learn its vocalizations. Ring dove males were exposed to conspecific vocalizations (coos), to heterospecific vocalizations (zebra finch song) or they were kept in silence. IEG expression was investigated by means of immunocytochemical analysis of the distribution of the ZENK protein product Zenk. In all three groups there was Zenk expression in several forebrain regions including the NCM and the CMM, similar to earlier findings in song-learning species. Quantitative analysis of the NCM, the CMM, and the hippocampus revealed significantly greater Zenk expression in birds exposed to conspecific vocalizations than in birds kept in silence, in the CMM only. These results show that there is substantial Zenk expression in the forebrain of a non-song-learning bird, comparable to that in avian song-learning taxa. These findings suggest that there is IEG expression specific to conspecific auditory stimulation in the CMM in both songbirds and non-songbirds.

Oral Administration of Melatonin to Old Ring Doves (Streptopelia risoria) Increases Plasma Levels of Melatonin and Heterophil Phagocytic Activity

We analyzed the effect of oral melatonin (23 microg/0.1 ml/animal/d; 1 h before dark on 12 consecutive days) in old birds, in natural photoperiods, on the hormone's plasma levels, and phagocytosis. Blood collections were performed daily at 2:00 am and 4:00 pm until 5 days after the treatment. From day 1, the melatonin levels were significantly higher than basal levels at both times. Values at 2:00 am were significantly higher than the 4:00 pm values. After treatment, the melatonin levels declined, returning from day 14 to basal values at both hours. At 2:00 am, phagocytosis was significantly greater than that obtained at 4:00 pm and greater than basal values. The 4:00 pm values were only significantly greater than basal on days 6 and 8, parallel to a decline in superoxide anion levels, which were lowest at 2:00 am. Melatonin administered to old ring doves increases the differences between nocturnal and diurnal plasma levels, and, in parallel, increases phagocytosis and reduces superoxide radical levels in heterophils.

Vocal tract filtering and the “coo” of doves

Ring doves (Streptopelia risoria) produce a "coo" vocalization that is essentially a pure-tone sound at a frequency of about 600 Hz and with a duration of about 1.5 s. While making this vocalization, the dove inflates the upper part of its esophagus to form a thin-walled sac structure that radiates sound to the surroundings. It is a reasonable assumption that the combined influence of the trachea, glottis and inflated upper esophagus acts as an effective band-pass filter to eliminate higher harmonics generated by the vibrating syringeal valve. Calculations reported here indicate that this is indeed the case. The tracheal tube, terminated by a glottal constriction, is the initial resonant structure, and subsequent resonant filtering takes place through the action of the inflated esophageal sac. The inflated esophagus proves to be a more efficient sound radiating mechanism than an open beak. The action of this sac is only moderately affected by the degree of inflation, although an uninflated esophagus is inactive as a sound radiator. These conclusions are supported by measurements and observations that have been reported in a companion paper.

Inflation of the esophagus and vocal tract filtering in ring doves

Ring doves vocalize with their beaks and nostrils closed, exhaling into inflatable chambers in the head and neck region. The source sound produced at the syrinx contains a fundamental frequency with prominent second and third harmonic overtones, but these harmonics are filtered out of the emitted signal. We show by cineradiography that the upper esophagus, oral and nasal cavities collect the expired air during vocalization and that the inflated esophagus becomes part of the suprasyringeal vocal tract. The level of the second and third harmonics, relative to the fundamental frequency (f(0)), is reduced in the esophagus and emitted vocalization compared with in the trachea, although these harmonics are still considerably higher in the esophagus than in the emitted signal. When the esophagus is prevented from fully inflating, there is a pronounced increase in the level of higher harmonics in the emitted vocalization. Our data suggest that the trachea and esophagus act in series as acoustically separate compartments attenuating harmonics by different mechanisms. We hypothesize that the trachea behaves as a tube closed at the syringeal end and with a variable, restricted opening at the glottal end that lowers the tracheal first resonance to match the f(0) of the coo. The inflated esophagus may function as a Helmholtz resonator in which the elastic walls form the vibrating mass. Such a resonator could support the f(0) over a range of inflated volumes.

Hormonal regulation of brain circuits mediating male sexual behavior in birds

Male sexual behavior in both field and laboratory settings has been studied in birds since the 19th century. Birds are valuable for the investigation of the neuroendocrine mechanisms of sexual behavior, because their behavior can be studied in the context of a large amount of field data, well-defined neural circuits related to reproductive behavior have been described, and the avian neuroendocrine system exhibits many examples of marked plasticity. As is the case in other taxa, male sexual behavior in birds can be usefully divided into an appetitive phase consisting of variable behaviors (typically searching and courtship) that allow an individual to converge on a functional outcome, copulation (consummatory phase). Based primarily on experimental studies in ring doves and Japanese quail, it has been shown that testosterone of gonadal origin plays an important role in the activation of both of these aspects of male sexual behavior. Furthermore, the conversion of androgens, such as testosterone, in the brain to estrogens, such as 17β-estradiol, is essential for the full expression of male-typical behaviors. The localization of sex steroid receptors and the enzyme aromatase in the brain, along with lesion, hormone implant and immediate early gene expression studies, has identified many neural sites related to the control of male behavior. The preoptic area (POA) is a key site for the integration of sensory inputs and the initiation of motor outputs. Furthermore, prominent connections between the POA and the periaqueductal gray (PAG) form a node that is regulated by steroid hormones, receive sensory inputs and send efferent projections to the brainstem and spinal cord that activate male sexual behaviors. The sensory inputs regulating avian male sexual responses, in contrast to most mammalian species, are primarily visual and auditory, so a future challenge will be to identify how these senses impinge on the POA–PAG circuit. Similarly, most avian species do not have an intromittent organ, so the projections from the POA–PAG to the brainstem and spinal cord that control sexual reflexes will be of particular interest to contrast with the well characterized rodent system. With this knowledge, general principles about the organization of male sexual circuits can be elucidated, and comparative studies relating known species variation in avian male sexual behaviors to variation in neural systems can be pursued.

Prolactin-induced parental hyperphagia in ring doves: are glucocorticoids involved?

Hyperphagia is a prominent component of the parental behavior repertoire in male and female ring doves and is necessary in order for parents to successfully provision their growing young. Although previous studies implicate both prolactin and the endogenous glucocorticoid, corticosterone, in parental hyperphagia, the functional interactions between these two hormones in regulating changes in feeding activity have not been characterized. These studies examined the possibility that prolactin's orexigenic effects are mediated through the increased secretion of corticosterone. Twice-daily intracerebroventricular (icv) injection of prolactin increased plasma corticosterone concentration in non-breeding doves of both sexes, with males exhibiting more pronounced effects than females. To further test the importance of glucocorticoid signaling in prolactin-induced feeding responses, changes in food intake were investigated in icv prolactin-treated, non-breeding doves following icv infusion of the glucocorticoid receptor antagonist RU38486 or propylene glycol vehicle. No attenuation of prolactin-induced hyperphagia was observed in either sex following co-administration of RU38486 at a dose shown previously to block dexamethasone-induced feeding in doves. These findings suggest that elevated corticosterone titers in blood may contribute to the hyperphagia observed in response to prolactin, but corticosterone signaling through a mammalian-type glucocorticoid receptor is not essential.

Temporal specificity of extinction in autoshaping.

Three experiments investigated the effects of varying the conditioned stimulus (CS) duration between training and extinction. Ring doves (Streptopelia risoria) were autoshaped on a fixed CS-unconditioned stimulus (US) interval and extinguished with CS presentations that were longer, shorter, or the same as the training duration. During a subsequent test session, the training CS duration was reintroduced. Results suggest that the cessation of responding during an extinction session is controlled by generalization of excitation between the training and extinction CSs and by the number of nonreinforced CS presentations. Transfer of extinction to the training CS is controlled by the similarity between the extinction and training CSs. Extinction learning is temporally specific.

Phagocytosis of Candida albicans and superoxide anion Levels in ring dove (Streptopelia risoria) heterophils: effect of melatonin.

We observed in previous studies on avian heterophils that incubation with either physiological or pharmacological concentrations of the neurohormone melatonin increased the phagocytosis of inert particles (latex beads), and also provoked a decline in superoxide anion levels of those phagocytes. In the present study, we wanted to corroborate whether melatonin acts on the oxidative metabolism that accompanies the respiratory burst during phagocytosis by inducing a more effective phagocytic activity at the same time as exerting an antioxidant effect to eliminate and/or scavenge the free radicals left over after the destruction of the foreign material. To this end, we evaluated the ingestion and destruction of Candida albicans (live particles) by ring dove (Streptopelia risoria) heterophils after different times of incubation (30 and 60 min) with physiological concentrations of melatonin (50 pg/ml diurnal and 300 pg/ml nocturnal), as well as with a pharmacological concentration 23 x 106 pg/ml (100 micro m) of the hormone. In parallel, using the same times of incubation, we evaluated the oxidative metabolism by determining the superoxide anion levels (O2-.). The results show that melatonin, at all the times and concentrations studied, increases both the phagocytosis index (number of C. albicans phagocytosed by 100 heterophils) and the candidicide power (percentage of C. albicans killed of those ingested by 100 heterophils). The effect was dose-dependent. With respect to the oxidative metabolism accompanying the digestion and destruction, there was a decline in superoxide anion levels after incubation with all of the concentrations of the hormone studied. The effect was dose-dependent and most pronounced at 60 min. These results thus corroborate the proposal that melatonin enhances the phagocytic function at the same time as neutralizing the oxidative stress derived from this immune function.

Changes in agouti-related peptide during the ring dove breeding cycle in relation to prolactin and parental hyperphagia.

Ring doves (Streptopelia risoria) exhibit marked increases in food consumption and decreases in body weight when they are provisioning their young. This study examined changes in hypothalamic immunostaining for agouti-related peptide (AGRP), an endogenous melanocortin receptor antagonist and appetite stimulant, during the ring dove breeding cycle. Because prolactin is orexigenic in doves, and is elevated in blood at the onset of parental hyperphagia, we also explored the possibility that prolactin-induced hyperphagia is associated with AGRP changes. The numbers of AGRP-immunoreactive (ir) cells within the tuberal hypothalamus were maximal during the prelaying period of the breeding cycle but decreased dramatically during early incubation. AGRP-ir cell numbers began to increase again during late incubation and reached a peak during the early and late posthatching stages. Because posthatching elevations in AGRP-ir were temporally associated with marked elevations in parental food intake, and because AGRP is orexigenic in doves, these findings suggest that increased AGRP activity in the dove tuberal hypothalamus may contribute to parental hyperphagia. Rising prolactin secretion during late incubation and early posthatching may initiate this increase in AGRP-ir, since intracerebroventricular administration of prolactin significantly elevated food intake and AGRP-ir cell numbers in the tuberal hypothalamus of nonbreeding doves. Prolactin-induced elevations in AGRP-ir cell numbers persisted when the confounding effects of weight gain that accompany prolactin-induced hyperphagia in nonbreeding doves were eliminated by a food restriction procedure, thereby suggesting that prolactin can directly influence AGRP activity under neutral energy state conditions.

Wing kinematics of avian flight across speeds

To test whether wing shape affects the kinematics of wing motion during bird flight, we recorded high‐speed video (250 Hz) of four species flying in a variable‐speed wind tunnel. The birds flew at intervals of 2 m s−1, ranging from 1 m s−1 up to their respective maximum flight speed, which varied from 14 to 17 m s−1 depending on the species. Kinematic data obtained from two synchronized, high‐speed video cameras were analyzed using 3D reconstruction. Three species with relatively pointed, high‐aspect ratio wings changed wingbeat styles according to flight speed (budgerigar, Melopsittacus undulatus; cockatiel, Nymphicus hollandicus; ringed turtle dove, Streptopelia risoria). These species used a wing‐tip reversal upstroke, characterized by supination of the distal wing at mid‐upstroke, at equivalent airspeeds ≤7 to 9 m s−1. In faster flight, they used a swept‐wing upstroke, without distal wing supination. At mid‐upstroke at any speed, wingspan in these species was greater than wrist span. In contrast, at all steady flight speeds, the black‐billed magpie Pica hudsonia with relatively broad, low‐aspect ratio wings, used a flexed‐wing, feathered upstroke in which wrist spans were equal to or greater than wingspans. Our results demonstrate that wing kinematics vary gradually as a function of flight speed, and that the patterns of variation are strongly influenced by external wing shape.

Sex steroid communication in the ring dove brain during courtship.

This review examines possible role of progesterone receptor (PR) and androgen receptor (AR) "cross-talk" in the expression of courtship behaviour in the ring dove (Streptopelia risoria). In doves, although androgen has been mostly associated with aggressive courtship behaviour and progesterone with the initiation of incubation, progesterone administration to courting birds terminates the aggressive component of courtship whilst having no effect on nesting behaviour. Recent results in doves have identified a high density of androgen receptor and progesterone receptor immunoreactivity (AR-ir and PR-ir) in the hypothalamus of both sexes in regions known to be directly involved in courtship and incubation behaviour. Nuclear AR-ir in courting birds is widespread throughout the brain. Nuclear PR-ir is only localized in discrete regions of the preoptic hypothalamus of both sexes. In the anterior and posterior hypothalamus of courting birds an increase number of AR-ir and PR-ir neurons colocalizes (70-90%) in the nucleus preopticus anterior (POA), nucleus preopticus medialis (POM), nucleus preopticus paraventricularis magnocellularis (PPM), nucleus hypothalami lateralis posterioris (PLH), and tuberal hypothalamus (Tu). A lower percentage of colocalization is seen in birds at other stages of the breeding cycle. The high percentage of AR-ir and PR-ir colocalization in the preoptic hypothalamus of courting doves supports previous reports involving progesterone acting in these brain regions to terminate the androgen-dependent aggressive courtship behaviour in male doves. The increase in PR-ir staining intensity in AR-ir neurons in courting birds suggests that this progesterone-dependent termination of aggressive courtship display in males occurs at the receptor level and may be orchestrated by central oestrogen.

Lack of changes in the courtship behaviour of male ring doves after testosterone treatment

The courtship behaviour of male birds depends on gonadal androgen. To date, however, there is little evidence that the expression and the features of courtship behaviours are proportional to the concentration of circulating androgen. The existence of such dose-dependent relationship is at the basis of several current theories of sexual selection. In the male ring dove (Streptopelia risoria) gonadal testosterone (T) controls courtship displays either directly (aggressive displays, i.e. bowing and chasing) or after conversion into oestradiol within the brain (nest-oriented displays). We studied whether the treatment of intact, sexually active males with T induces short-term changes in the fine structure of the bowing display and in the frequency of the courtship displays. Because nestsoliciting depends on brain oestrogen formation, we also measured the activity of the oestrogen-synthesising enzyme aromatase in the brain areas controlling courtship. The structure of the bowing display and the pattern of the courtship were not affected by either of two T treatments that elevated circulating androgen within the physiological range. Males implanted with the higher dose of T had higher aromatase activity in the anterior hypothalamus. Our results indicate that T modulates brain aromatase in intact males and suggest a threshold model of action of steroid hormones on the structure of courtship behaviour.

Season-related changes in circulating androgen, brain aromatase, and perch-calling in male ring doves

The perch-call of ring doves ( Streptopelia risoria) is related to territorial defence and mate attraction. Perch-calls are sexually dimorphic and individually different in structure. The expression of perch-call is androgen-dependent and is controlled by the action of testosterone on the preoptic-hypothalamic areas. However, it is not known whether the acoustic features of the call vary with the reproductive condition. We studied plasma androgen levels, brain steroid metabolism, and perch-calling of male ring doves kept in winter-like (low temperature, short days) or spring-like (mild temperature, long day) conditions. Circulating levels of androgen were higher in males kept on spring-like condition. Spring males spent more time in perch-calling than winter males. However, a detailed analysis of the call structure revealed no difference for any time or frequency parameter between groups. This work shows that in ring doves season-dependent variations in the circulating levels of androgen are correlated with differences in the amount of time males spend in perch-calling. In addition, the study suggests that in adult males the acoustic structure of the perch-calls is not influenced by season-related changes in androgen levels.