Gap detection in the starling (Sturnus vulgaris)

Summary Gap-detection thresholds of single units were determined from auditory forebrain neurons of the awake starling. Nine different response types were statistically defined from the discharge pattern to a 400 ms broadband noise stimulus. The gap stimuli consisted of two broadband noise bursts which were separated by a gap ranging from 0.4 to 204.8 ms duration. The median minimumdetectable gap for 121 out of 145 units that had a significant threshold 204.8ms was 12.8 ms; 20% of the neurons showed thresholds between 0.4 and 3.2 ms. The neurons of the nine response types differed significantly in their minimum-detectable gaps; neurons with phasic-tonic and phasic excitation exhibited the best (i.e. shortest) minimum-detectable gaps. The neurons of the three different recording areas (field L, NCM and HV) were significantly different in their minimumdetectable gaps; field L neurons showed the best temporal resolution for gaps in broadband noise. Gap-detection thresholds are compared with psychophysical thresholds determined with the same stimuli and the relevance of forebrain units for temporal resolution is discussed.Abbreviations CS control stimulus - HV hyperstriatum ventrale - HVc hyperstriatum ventrale pars caudalis - NB noise burst - NCM neostriatum caudale pars medialis - NS noise stimulus - SGS standard gap series - TW time window     

Gap detection in the European starling (Sturnus vulgaris)

Summary Gap-detection thresholds were determined for single units in the cochlear ganglion and in auditory nerve fibres of the starling from responses to two broad-band noise bursts separated by a temporal gap of between 0.4 and 204.8 ms. All 35 units showed a threshold within the range of gap sizes tested. The median minimum-detectable gap was 12.8 ms with the minimum being 1.6 ms. A multiple regression analysis revealed that the size of the minimum-detectable gap was not significantly correlated with the neuron's CF, with its sharpness of tuning as given by its bandwidth 10 dB above threshold, or with its Q_10dB value. Only the level of stimulation above the neuron's threshold showed a significant negative correlation with the size of the minimum-detectable gap. These results are discussed with respect to theoretical considerations of limits posed on temporal resolution by the characteristics of peripheral filters. These findings are also discussed in the context of the coding of gaps at different levels of the starling's auditory system and in relation to psychoacoustic results in the starling on gap detection and time resolution described by temporal modulation transfer functions.     

Azimuthal sound localization in the European starling (Sturnus vulgaris)

Summary The physical measurements reported here test whether the European starling (Sturnus vulgaris) evaluates the azimuth direction of a sound source with a peripheral auditory system composed of two acoustically coupled pressure-difference receivers (1) or of two decoupled pressure receivers (2).A directional pattern of sound intensity in the freefield was measured at the entrance of the auditory meatus using a probe microphone, and at the tympanum using laser vibrometry. The maximum differences in the soundpressure level measured with the microphone between various speaker positions and the frontal speaker position were 2.4 dB at 1 and 2 kHz, 7.3 dB at 4 kHz, 9.2 dB at 6 kHz, and 10.9 dB at 8 kHz. The directional amplitude pattern measured by laser vibrometry did not differ from that measured with the microphone. Neither did the directional pattern of travel times to the ear. Measurements of the amplitude and phase transfer function of the starling's interaural pathway using a closed sound system were in accord with the results of the free-field measurements.In conclusion, although some sound transmission via the interaural canal occurred, the present experiments support the hypothesis 2 above that the starling's peripheral auditory system is best described as consisting of two functionally decoupled pressure receivers.Abbreviations CM cochlear microphonics - ITD interaural time difference - IID interaural intensity difference - MRA minimum resolvable angle - dB SPL sound-pressure level (re 0.00002 Pa)     

Innervation of the parathyroid in the european starling (Sturnus vulgaris)

Anatomical studies were conducted to characterize the source, type, and distribution of parathyroid gland innervation in European starlings. Denervation experiments demonstrated that the parathyroid glands and adjacent carotid bodies are innervated by nerve fibers originating in the nodose ganglion of the vagus nerve. In the parathyroid parenchyma, these fibers terminate adjacent to chief cells or near vascular smooth muscle. Vagal fibers also form synapses with catecholamine-containing glomus cells of the carotid body. Blood that first perfuses the carotid body subsequently perfuses the parathyroid parenchyma. These observations suggest that vagal innervation may influence parathyroid function in starlings either through direct chief cell innervation or through alteration of vascular perfusion. A neurohemal relationship also may exist between the carotid body and parathyroids.     

Metabolic costs of egg production in the European starling (Sturnus vulgaris)

The energy cost of egg production in passerine birds has typically been estimated to be 45%-60% of basal metabolic rate (BMR), but this is based on theoretical models using data on energy content of eggs and reproductive tissue; there are still very few empirical data on egg production costs. In this study, we directly measured resting metabolic rate (RMR) in egg-laying female European starlings (Sturnus vulgaris) over 3 yr. We compared these data with RMR of nonbreeding and chick-rearing birds and with estimated energy expenditure generated from a typical energy content model by using empirically derived data from body composition analysis for this species. We found marked variation in RMR between years and between reproductive stages, which complicates comparisons among breeding stages for the assessment of relative egg production costs. On the basis of this method, RMR during egg laying varied from +74% to -13% of nonbreeding RMR and from +20% to -7% of chick-rearing RMR. We therefore used an alternate approach: measuring changes in RMR through the complete cycle of follicle development and ovulation. The increase in RMR from the beginning of prelaying to the six-follicle stage (before first ovulation) when birds have a complete developing follicle hierarchy was 22.4%. This value is still much lower than that estimated from our energy content model. We discuss conceptual problems associated with the theoretical energy content approach but also suggest, on the basis of earlier work done in our lab, that the measured increase in RMR might still underestimate the actual cost of egg production if birds reallocate energy between different physiological systems.     

Foraging rate versus sociality in the starling Sturnus vulgaris

It is well established that social conditions often modify foraging behaviour, but the theoretical interpretation of the changes produced is not straightforward. Changes may be due to alterations of the foraging currency (the mathematical expression that behaviour maximizes) and/or of the available resources. An example of the latter is when both solitary and social foragers maximize rates of gain over time, but competition alters the behaviour required to achieve this, as assumed by ideal free distribution models. Here we examine this problem using captive starlings Sturnus vulgaris. Subjects had access to two depleting patches that replenished whenever the alternative patch was visited. The theoretical rate-maximizing policy was the same across all treatments, and consisted of alternating between patches following a pattern that could be predicted using the marginal value theorem (MVT). There were three treatments that differed in the contents of an aviary adjacent to one of the two patches (called the 'social' patch). In the control treatment, the aviary was empty, in the social condition it contained a group of starlings, and in a non-specific stimulus control it contained a group of zebra finches. In the control condition both patches were used equally and behaviour was well predicted by the MVT. In the social condition, starlings foraged more slowly in the social than in the solitary patch. Further, foraging in the solitary patch was faster and in the social patch slower in the social condition than in the control condition. Although these changes are incompatible with overall rate maximization (gain rate decreased by about 24% by self-imposed changes), if the self-generated gain functions were used the MVT was a good predictor of patch exploitation under all conditions. We discuss the complexities of nesting optimal foraging models in more comprehensive theoretical accounts of behaviour integrating functional and mechanistic perspectives.     

The functional anatomy of the shoulder in the European starling (Sturnus vulgaris)

The excursions of wing elements and the activity of eleven shoulder muscles were studied by cineradiography and electromyography in European starlings (Sturnus vulgaris) flying in a wind tunnel at speeds of 9–20 m s^?1. At the beginning of downstroke the humerus is elevated 80–90° above horizontal, and both elbow and wrist are extended to 90° or less. During downstroke, protraction of the humerus (55°) remains constant; elbow and wrist are maximally extended (120° and 160°, respectively) as the humerus passes through a horizontal orientation. During the downstroke-upstroke transition humeral depression ceases (at about 20° below horizontal) and the humerus begins to retract. However, depression of the distal wing continues by rotation of the humerus and adduction of the carpometacarpus. Humeral retraction (to within about 30° of the body axis) is completed early in upstroke, accompanied by flexion of the elbow and carpometacarpus. Thereafter the humerus begins to protract as elevation continues. At mid-upstroke a rapid counterrotation of the humerus reorients the ventral surface of the wing to face laterad; extension of the elbow and carpometacarpus are initiated sequentially. The upstroke-downstroke transition is characterized by further extension of the elbow and carpometacarpus, and the completion of humeral protraction. Patterns of electromyographic activity primarily coincide with the transitional phases of the wingbeat cycle rather than being confined to downstroke or upstroke. Thus, the major downstroke muscles (pectoralis, coracobrachialis caudalis, sternocoracoideus, subscapularis, and humerotriceps) are activated in late upstroke to decelerate, extend, and reaccelerate the wing for the subsequent downstroke; electromyographic activity ends well before the downstroke is completed. Similarly, the upstroke muscles (supracoracoideus, deltoideus major) are activated in late downstroke to decelerate and then reaccelerate the wing into the upstroke; these muscles are deactivated by mid-upstroke. Only two muscles (scapulohumeralis caudalis, scapulotriceps) exhibit electromyographic activity exclusively during the downstroke. Starlings exhibit a functional partitioning of the two heads of the triceps (the humerotriceps acts with the pectoralis group, and does not overlap with the scapulotriceps). The biphasic pattern of the biceps brachii appears to correspond to this partitioning.     

Intraspecific nest parasitism in the European starling Sturnus vulgaris

Biochemical genetic markers were used along with conventional methods (abnormal laying sequence/clutch size, unusual egg shape/pigmentation) to identify intraspecific nest parasitism at two British nestbox colonies of the European starling. Between 11 and 37% of first clutches were parasitized during 1977–1979. Parasitic females probably comprised all of the following categories: (1) paired females contesting a nestbox occupied by another pair; (2) previously paired females who had laid a clutch but had been unsuccessful; (3) unpaired females who had copulated with males that already had a mate and nest site; and (4) ‘professional’ nest parasites who distributed at lest some of their eggs in one or more nests other than their own. Although parasitized nests had higher clutch sizes, parasitism led to fewer host young fledging per egg laid, mainly through the eviction of eggs and subsequent nest desertion. Number of parasitic young fledged per egg laid was highest when eggs were laid synchronously with the host, when host clutches were larger, or a smaller number of parasite eggs were added to a nest, thus favouring parasites that distribute their eggs amongst a number of nests. A greater pressure on nest sites may have accounted for the higher levels of parasitism at the Aberdeen colony and for the greater number of parasite eggs laid in a nest. Although most parasitic female starlings appeared to be much less successful than non-parasitic ones, nest parasitism in the starling might evolve directly when one or more of the following advantages are present. (1) There are no constraints on the number of eggs a female may lay but there are constraints on the number of young she may feed adequately. (2) Female survival is increased by having fewer or no eggs/young to care for. (3) Current feeding conditions favour the survival of more young than would be produced by the most common clutch size. Intraspecific nest parasitism is considered to be a first stage in the evolution of interspecific nest parasitism.     

A dynamic model of food allocation to starling (Sturnus vulgaris) nestlings

We present a dynamic model of food allocation to starling nestlings.Based on the premise that current reproductive activities mayhave detrimental effects on future breeding episodes, the modelis elaborated around the concept of a state variable. In thisimplementation, the state variable represents body mass. A numberof activities can be followed during the breeding season, andwe measured the value of these actions in terms of the contributionof the action to the fitness of the animal at the end of thenestling period. Using the final states reached by parents andyoung, we determined fitness by summing current breeding productionand future reproductive potential. We used backward inductionto find for each combination of states and nestling age thebehavioral options that maximize fitness. Subsequently, forwardsimulations were used to find average trajectories across timeof the following variables: parental and offspring states, foragingtime, and allocation of energy. We also investigated the effectsof brood size and harvest. Physiological parameters, necessaryto translate foraging decisions into body mass changes, aretaken from studies on starlings (Sturnus vulgaris). The resultsappear to closely fit available evidence for this species. Thediscussion relates basic findings of the model to previous resultsof static and deterministic approaches. Limits to the usefulnessof the model are discussed. [Behav Ecol 1991; 2: 21-37]     

Isolation and characterization of polymorphic microsatellites isolated from the spotless starling (Sturnus unicolor) and cross‐species amplification in the European starling (Sturnus vulgaris)

Spotless and European starlings (Sturnus unicolor and Sturnus vulgaris) have attracted attention from researchers interested in sexual selection, evolution of parental care and reproductive strategies. Both species show high levels of intraspecific nest parasitism and extra‐pair paternity, but research in this area is hampered by a lack of molecular markers specific for these species. Here, we describe a set of primers for nine microsatellite loci in spotless starlings, eight of which are highly polymorphic (7.37 alleles in average). These microsatellites are also polymorphic in European starlings (6.75 alleles in average).     

Ahemeral light regimens test the photoperiodic threshold of the european starling(Sturnus vulgaris)

Male European starlings(Sturnus vulgaris) were held for three consecutive photoperiod oscillations (ahemeral years) composed of 30-h day lengths, i.e., the daily light and dark each lasted three hours longer than under the natural daily photoperiod at latitude 38°N. These starlings had no gonad metamorphosis during the 45 actual months necessary to complete the three ahemeral photoperiod oscillations; nor did subsequent exposure to continuous illumination elicit gonad response. It is concluded that the daily duration of light and darkness (although certainly operant in controlling starling sexual cycles under temperate-zone photoperiod oscillations) is not the critical factor establishing a sexual cycle under the ahemeral regimen. Rather, it appears that this species must experience a daily duration of light of 12 hours or less (a definitive photoperiodic threshold) before photo-induction of a sexual cycle is possible.Presented at the Eighth International Congress of Biometeorology, 9–14 September 1979, Shefayim, Israel.     

Developmental stress, social rank and song complexity in the European starling (Sturnus vulgaris)

Bird song is a sexually selected trait and females have been shown to prefer males that sing more complex songs. However, for repertoire size to be an honest signal of male quality it must be associated with some form of cost. This experiment investigates the effects of food restriction and social status during development on song complexity in the European starling (Sturnus vulgaris). Birds that experienced an unpredictable food supply early in life produced a significantly smaller repertoire of song phrases than those with a constant food supply. Social status during development was also significantly correlated with repertoire size, with dominant birds producing more phrase types. This study therefore provides novel evidence that social as well as nutritional history may be important in shaping the song signal in this species.     

The breeding biology of the starling Sturnus vulgaris in northern Finland

The breeding biology of the starling Sturnus vulgaris was studied in three populations in northern Finland, with additional reference localities in nearby areas. Results from a total of 329 nests are presented from years 1962-1976.
The onset of laying in the Oulu area took place on 5 May, the median date being 10 May, about one week later than in southern Finland and 2–3 weeks later than in central Europe. The clutch-size was 5.29 ± 0.05 in the first nests and 5.06 ± 0.26 in later ones. Incubation time was 12.1 ± 0.6 d. An average of 3.31 ± 0.10 fledglings was produced per pair, 3.69 ± 0.08 in succesful clutches, which is significantly less than usually reported from central Europe. Most of the losses were due to starvation of the young.
No second broods were observed in the Oulu area, and repeated nesting was not common, probably because of the general late start of laying.
The reasons for the recent decline in several Finnish starling populations do not lie in the production capacity but must be sought for elsewhere.     

Female presence affects male behavior and testosterone levels in the European starling (Sturnus vulgaris)

In this study, we confronted individually housed male European starlings (Sturnus vulgaris) with a female conspecific for 60 min to study the consequences on behavior and plasma testosterone (T) concentrations. Control males experienced a similar procedure, the only difference being that they were tested in the absence of a female. Female presence significantly affected both behavior and plasma T levels of male starlings. Experimental males spent significantly more time singing in the nest box and flew significantly more into the nest box with green nesting material during female presentations than during control periods. Control males never showed these mate attraction behaviors. In total 5 of the 16 experimental males did not respond behaviorally to the female stimulus bird (NR males). In contrast to T levels of control males, plasma T concentrations of both experimental males that did respond to the female (R males) and of NR males (which only perceived the female stimulus) were positively influenced by female presentation. The time spent singing in the nest box by experimental males (R and NR males combined) during female presence tended to be positively correlated with changes in plasma T levels. Finally, before introduction of a female, plasma T levels of R males were significantly higher than those of NR males indicating that individually housed males respond to the presence of a female conspecific by increasing their mate attraction behaviors only when a threshold plasma T concentration has been reached.     

Azimuthal sound localization in the European starling (Sturnus vulgaris): II. Psychophysical results

Small songbirds have a difficult analysis problem: their head is small compared to the wavelengths of sounds used for communication providing only small interaural time and level differences. Klump and Larsen (1992) measured the physical binaural cues in the European starling (Sturnus vulgaris) that allow the comparison of acoustical cues and perception. We determined the starling’s minimum audible angle (MAA) in an operant Go/NoGo procedure for different spectral and temporal stimulus conditions. The MAA for broadband noise with closed-loop localization reached 17°, while the starling’s MAA for open-loop localization of broadband noise reached 29°. No substantial difference between open-loop and closed-loop localization was found in 2 kHz pure tones. The closed-loop MAA improved from 26° to 19° with an increase in pure tone frequency from 1 to 4 kHz. This finding is in line with the physical cues available. While the starlings can only make use of interaural time difference cues at lower frequencies (e.g., 1 and 2 kHz), additional interaural level difference cues become available at higher frequencies (e.g., 4 kHz or higher, Klump and Larsen 1992). An improvement of the starling’s MAA with an increasing number of standard stimulus presentations prior to the test stimulus has important implications for determining relative (MAA) localization thresholds.     

Lesions to the medial preoptic area affect singing in the male European starling (Sturnus vulgaris)

The aromatization of testosterone (T) in the medial preoptic nucleus (POM) is known to regulate male courtship and sexual behaviors expressed prior to, and in anticipation of, copulation. Singing in male European starlings is used to attract mates prior to physical sexual contact, suggesting that the POM might be involved. The present study was performed to examine the effects of lesions targeting the POM on singing and courtship behavior in reproductively active male starlings. A significant decrease in song output and the gathering of green nest materials was observed in males with lesions to the POM compared to males with damage to brain areas outside of the POM. Lesions did not affect a male's tendency to remain near a female or to occupy a nestbox, suggesting that the effects of POM lesions were specific to courtship behaviors. Behavioral differences were not related to testis mass or volume, and GnRH immunoreactivity was observed within the hypothalamus and median eminence for each male, suggesting that the effects of POM lesions were related specifically to POM involvement in song expression rather than to a disruption of the GnRH axis. These results suggest a general role for the POM in the expression of behaviors related to sexual arousal or anticipation, including song.     

Allozyme polymorphism and interspecific relationships in the Common starling (Sturnus vulgaris) and Spotless starling (S. unicolor) (Aves: Sturnidae)

This paper reports a genetic study of Sturnus vulgaris and S. unicolor (Sturnidae), two similar bird species which have recently become sympatric in north-eastern Spain. Seven enzyme systems (alcohol dehydrogenase, esterase, glutamate dehydrogenase, glucose-6-phosphate dehydrogenase, leucin aminopeptidase, L-lactate dehydrogenase and superoxide dismutase) were analyzed, resolving 22 presumed genetic loci with 28 alleles; 27 of them were found in both species. Gene flow was estimated to be number of migrants per generation (Nm) = 6, and genetic identities among populations were high, ranging from I = 0.96 to I = 0.99. On the whole, the results demonstrate that these Sturnus populations share a gene pool, showing slight genetic differences between the two starling speices.     

Song as an honest signal of past developmental stress in the European starling (Sturnus vulgaris)

Bird song is a sexually selected male trait where females select males on the basis of song quality. It has recently been suggested that the quality of the adult male song may be determined by nutritional stress during early development. Here, we test the 'nutritional-stress hypothesis' using the complex song of the European starling. Fledgling starlings were kept under experimental treatment (unpredictable short-term food deprivations) or control conditions (ad libitum food supply), for three months immediately after independence. We measured their physiological and immune responses during the treatment and recorded song production during the following spring. Birds in the experimental group showed increased mass during the treatment and also a significantly suppressed humoral response compared with birds in the control group. There was no difference between the groups in the cell-mediated response. Next spring, males in the experimental group spent less time singing, sang fewer song bouts, took longer to start singing and also sang significantly shorter song bouts. These data support the hypothesis that both the quality and quantity of song produced by individual birds reflect past developmental stress. The results also suggest the 'nutritional-stress hypothesis' is best considered as a more general 'developmental-stress hypothesis'.     

Dynamics of Campylobacter colonization of a natural host, Sturnus vulgaris (European starling)

Wild European Starlings ( Sturnus vulgaris ) shed Campylobacter at high rates, suggesting that they may be a source of human and farm animal infection. A survey of Campylobacter shedding of 957 wild starlings was undertaken by culture of faecal specimens and genetic analysis of the campylobacters isolated: shedding rates were 30.6% for Campylobacter jejuni , 0.6% for C. coli and 6.3% for C. lari. Genotyping by multilocus sequence typing (MLST) and antigen sequence typing established that these bacteria were distinct from poultry or human disease isolates with the ST-177 and ST-682 clonal complexes possibly representing starling-adapted genotypes. There was seasonal variation in both shedding rate and genotypic diversity, both exhibiting a maximum during the late spring/early summer. Host age also affected Campylobacter shedding, which was higher in younger birds, and turnover was rapid with no evidence of cross-immunity among Campylobacter species or genotypes. In nestlings, C. jejuni shedding was evident from 9 days of age but siblings were not readily co-infected. The dynamics of Campylobacter infection of starlings differed from that observed in commercial poultry and consequently there was no evidence that wild starlings represent a major source of Campylobacter infections of food animals or humans.