Effects of begging on growth rates of nestling chicks

We investigated whether an increase in begging levels delaysgrowth of chicks. In experiment 1, we hand-reared nine pairsof ring dove squabs, divided into a control and a begging group.All squabs received similar amounts of food, but those in thebegging group had to beg for a prolonged period in order tobe fed, while squabs in the control group received food withoutbegging. Squabs stopped responding to the treatment after 10days and, at that time, there was no effect of induced beggingon their body mass. In experiment 2, we hand-reared 27 pairsof magpie chicks for 3 days. The design of experiment 2 wassimilar to that of experiment 1. Daily food intake and beggingaffected growth rates. On average, chicks in the begging groupgrew 0.8 g/day less than control chicks, which represents adecrease of 8.15% in growth rate. Because growth is usuallypositively associated with expected fitness, this demonstratesthat begging is a costly behavior, an assumption routinelymade in models of begging behavior.     

Yolk androgens and begging behaviour in black-headed gull chicks: an experimental field study

Black-headed gulls, Larus ridibundus, produce clutches of three eggs, which contain high levels of maternal androgens in the yolk. These levels increase with laying order and the eggs hatch asynchronously. Experiments have supported the hypothesis that this within-clutch variation in maternal androgens mitigates the disadvantage of last-hatched chicks in sibling competition, by enhancing embryonic development and early posthatching growth. We hypothesized that these effects come about by the stimulating effects of maternal androgens on begging behaviour and competitive ability. In the field, we injected first-laid eggs of a clutch (which have a low androgen level) with either an androgen solution (T eggs) or vehicle (Oil eggs). We then created pairs of chicks hatched from Oil and T eggs, matched for egg mass and hatching date. Parent-chick interactions were recorded from observation hides. Chicks from T eggs hatched almost half a day sooner than those from Oil eggs. Furthermore, chicks from T eggs were more active during the first week after hatching, were more often the first to react to the approaching parent, begged more frequently, and obtained the larger share of food. We conclude that the enhancing effect of yolk androgens on growth in this species arises at least partly through androgen-mediated effects on the chicks' behaviour. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.      

The hormonal control of begging and early aggressive behavior: experiments in black-headed gull chicks

The hormonal control of begging and sibling competition is largely unknown, but recent evidence suggests a role for steroid hormones. We tested the influence of the aromatizable androgen testosterone (T), the non-aromatizable androgen 5alpha-dihydrotestosterone (DHT), and 17beta-estradiol (E) on both begging behavior and aggressive behavior in black-headed gull chicks (Larus ridibundus). Chicks of this species have a conspicuous begging display, while their frequently performed early aggressive behavior is facilitated by testosterone and important for territorial defense. Hormone treatment was applied by implants between days 6 and 16 after hatching. Behavior was tested by means of standard stimulus tests. The results were validated in a second experiment under semi-natural conditions. Begging was suppressed by T and DHT and not affected by E. Aggressive Pecking was strongly facilitated by T. The erect threat posture, characteristic for older chicks, was facilitated by T, DHT, and E and the nest-oriented threat display, typical for young chicks, only by T and DHT. Growth was suppressed in the T group. The results indicate that androgen production, needed for territorial defense, has costs in terms of a suppression of begging and growth. It is discussed to what extent older chicks may avoid these costs by converting testosterone to estrogen and why pre-natal and post-natal exposure to androgens differ in their effect on begging behavior.     

Parental preference for red mouth of chicks in a songbird

Parental preferences during feeding and care-giving may select for ornamental traits in young, such as bright coloration. For chicks of coots, there is experimental evidence for this idea. We examined the hypothesis that bright yellow, orange and red mouths of chicks of songbirds have been favoured by feeding preferences in parents. In a field experiment, the orange–yellow mouths of great tit nestlings were dyed brightly red, and the feeding response of parents recorded. In nest boxes with extra daylight through a window, experimental chicks were on average given twice as much food (biomass) as control chicks (sham dyed). In normal nest boxes, the tendency was similar, but not significant. Thus, at least in good light, great tit parents prefer to feed young with red mouths, a preference for colourfulness that helps explain the evolution of bright gapes in chicks of songbirds (passerine birds).     

Effects of elevated yolk androgens on perinatal begging behavior in yellow-legged gull (Larus michahellis) chicks

Maternal hormones may represent an important pathway by which mothers can adaptively adjust offspring traits and performance to suit the prevailing environmental conditions. Earlier studies of birds have shown that egg androgens of maternal origin may enhance post-natal offspring 'begging' displays, functioning to solicit parental care. Here we investigate the effects of elevated egg androgen levels on the prenatal begging behavior of yellow-legged gull (Larus michahellis) chicks. At laying, we experimentally increased the concentration of yolk testosterone (T) within the natural range of variation, and, shortly before hatching, we compared the structural properties, rate, and loudness of vocalizations of embryos developing in T- and oil-injected (control) eggs. In addition, we compared the early post-hatch begging rate (measured as the pecking rate towards a dummy gull head) in chicks of the two experimental groups. We found that T embryos produced louder embryonic vocalizations than controls, whereas structural properties and the calling rate did not differ between T and control embryos. The post-hatch begging rate was unaffected by T treatment, but strongly decreased with increasing chick body mass, suggesting that intensity of the begging display was sensitive to chick state and may therefore reliably indicate the need of food in this species. Therefore, the results of this study show for the first time that prenatal T exposure modulates the quality of embryonic vocalizations, but are not in accordance with previous findings reporting increased post-hatching begging intensity following increased prenatal exposure to androgens.     

Response to begging calls by Zebra Finch parents: "first come, first served" rule may overcome a parental preference between chicks

In birds, parents may provide differential food provisioning among offspring according to their sex. Here, we test the hypothesis that events linked to the fine dynamics of begging behaviour could modulate parental preferences. After evaluating the preference related to chick sex for each parent of six Zebra Finch Taeniopygia guttata pairs, we studied the possible modifications of this preference when offspring begging was asynchronous. Our observations show that male parents follow a "first come, first served" rule, whereas females keep their initial choice. Although this study remains preliminary due to the sample size, it underlines the potential importance of investigating fine temporal features of begging behaviour to fully understand parents' provisioning strategies.     

Detectability matters: conspicuous nestling mouth colours make prey transfer easier for parents in a cavity nesting bird

An often underappreciated function of signals is to notify receivers of the presence and position of senders. The colours that ornament the mouthparts of nestling birds, for example, have been hypothesized to evolve via selective pressure generated by parents'' inability to efficiently detect and feed nestlings without such visually conspicuous targets. This proposed mechanism has primarily been evaluated with comparative studies and experimental tests for parental allocation bias, leaving untested the central assumption of this detectability hypothesis, that provisioning offspring is a visually challenging task for avian parents and conspicuous mouths help. To test this assumption, I manipulated the mouths of nestling house sparrows to appear minimally and maximally conspicuous, and quantified prey transfer difficulty as the total duration of a feeding event and the number of transfer attempts required. Prey transfer to inconspicuous nestlings was, as predicted, more difficult. While this suggests that detectability constraints could shape nestling mouth colour evolution, even minimally conspicuous nestlings were not prohibitively difficult for parents to feed, indicating that a more nuanced explanation for interspecific diversity in this trait is needed.     

Cooperative begging in banded mongoose pups

Vivid begging displays are common in species with parental care [1, 2]. They are usually seen as the way that rival offspring selfishly compete over parental investment [3], and individuals are expected to respond to the begging of rivals by increasing their own begging intensity [4, 5]. Here I show the opposite - that potential rivals gain direct benefits from begging by littermates, so that begging behavior becomes a collective enterprise, similar to other cooperative activities. I investigate begging in communally breeding banded mongooses (Mungos mungo), where each pup forms an exclusive relationship with a single helper (its "escort"), minimizing competition over food allocation. Escorts were influenced by the total signal emanating from a litter, so that pups who begged at low rates received more food as litter size increased. Focal pups increased their begging when litters were experimentally reduced or littermates were induced to beg at low rates, but they received food at similar rates and showed reduced weight gain - indicating that they were paying a higher cost for a similar reward. These results suggest that offspring can benefit from companions despite conflicts over the allocation of parental investment [6, 7]. Such benefits provide an explanation for observed variation in the expression of parent-offspring conflict.     

Urban begging and ethnic nepotism in Russia

Ethnic nepotism theory predicts that even in times of communal peace altruism is more pronounced within than between ethnic groups. The present study tested the hypothesis that altruism in the form of alms giving would be greater within than between ethnic groups, and greater between more closely related groups than between more distant groups. The three groups chosen for study were ethnic Russians, Moldavians, and Gypsies. Russians are genetically closer to Moldavians than to Gypsies. Observations were made of 128 ethnic Russian, 25 Moldavian, and 25 Gypsy beggars receiving gifts from ethnic Russians in Moscow trains. The Gypsies were mainly girls, contrary to the Russian sample. Multivariate analysis identified three main strategies: active, personified, and appeasing-undirected. Russian strategies were most variable. Gypsies presented strong charity releasers: 84% were children who played music and sang and showed appeasing-undirected behavior. The few adults were highly submissive or friendly. Nevertheless, their success was limited compared with that of ethnic Russians despite the latter’s demanding behavior and their being mostly mature or elderly persons. Moldavians received an intermediate amount of charity. The hypothesis was supported.     

Effect of rapid modulation of circulating plasma testosterone concentration on begging,aggressive behavior and competition for food in black-headed gull (Larus ridibundus) chicks

Sibling competition mediated by begging behavior is extremely common in avian species and recent studies have highlighted the role of endogenous testosterone in regulating such phenomenon. However, current literature depicts an inconsistent pattern in altricial vs. semi-precocial species, with stimulating versus inhibitory effects of the hormone respectively. This is possibly due to a difference in the methodology of hormone treatment (short-term moderate dose versus a long-term stronger elevation, respectively) between the studies performed so far. In this study, we induced short-term moderate peaks in plasma testosterone levels, as applied in altricial bird species, and assessed the effects of our manipulation on begging, competitive and aggressive behavior in black-headed gull (Larus ridibundus) chicks, a semi-precocial species. Our results suggest that, unlike in altricial songbirds, temporary increase of plasma testosterone concentration suppresses begging and enhances aggressiveness towards intruders. However, it also increases aggression and the chances of getting priority while scrambling with nest mates to gain access to food. Thus, the inconsistencies in the hormonal control of begging behavior observed between altricial vs. semi-precocial birds seem real and perhaps related to species differences in complexity of the display and the nature of competition. These may be elucidated by future comparative studies.     

Nestling testosterone controls begging behaviour in the pied flycatcher, Ficedula hypoleuca

Begging signals and endogenous testosterone (T) levels of young birds have been shown to be positively correlated. If T is causally involved in controlling the level of begging effort, an endocrine control mechanism could explain the evolution of begging as a costly signal reflecting need. We tested experimentally whether elevated circulating T levels enhanced begging behaviour in nestling pied flycatchers, Ficedula hypoleuca. A pilot study confirmed that nestling T levels could be elevated within a natural physiological range using an oral dose of T. After T-dosing, nestling begging behaviour was measured as: i) the duration of begging displays and ii) the maximum height of begging stretches. Our results show that nestling T levels were elevated at 90 min post dosing and that at this time point both measures of begging behaviour were performed more intensely by T-dosed nestlings than controls. Nestling begging displays in response to dosing varied between individuals, which in part was explained either by the date in the breeding season or nestling mass. The results of this study confirm the causal nature of T in controlling nestling begging signals and suggest that it may be part of the mechanism that controls begging behaviour in nestling birds.     

Tetrachromacy, oil droplets and bird plumage colours

There is a growing body of data on avian eyes, including measurements of visual pigment and oil droplet spectral absorption, and of receptor densities and their distributions across the retina. These data are sufficient to predict psychophysical colour discrimination thresholds for light-adapted eyes, and hence provide a basis for relating eye design to visual needs. We examine the advantages of coloured oil droplets, UV vision and tetrachromacy for discriminating a diverse set of avian plumage spectra under natural illumination. Discriminability is enhanced both by tetrachromacy and coloured oil droplets. Oil droplets may also improve colour constancy. Comparison of the performance of a pigeon's eye, where the shortest wavelength receptor peak is at 410 nm, with that of the passerine Leiothrix, where the ultraviolet-sensitive peak is at 365 nm, generally shows a small advantage to the latter, but this advantage depends critically on the noise level in the sensitivity mechanism and on the set of spectra being viewed. Accepted: 3 July 1998     

An immunological cost of begging in house sparrow nestlings

Parent–offspring conflict predicts that offspring should demand a greater parental investment than is optimal for their parents to deliver. This would escalate the level of offspring demand ad infinitum, but most of the models on the evolution of parent–offspring communication predict that begging must be costly, such costs limiting the escalation and defining an optimal level of begging. However, empirical evidence on this issue is mixed. A potential begging cost that remains to be accurately explored is a decrease in immunocompetence for offspring begging fiercely. This study experimentally analyses this cost in house sparrow (Passer domesticus) nestlings. A group of nestlings was forced to beg fiercely for a prolonged time while a control group begged at low levels, both groups receiving the same quantity of food. At the same time, the nestling response to an antigen (phytohaemagglutinin) was measured. Nestlings forced to beg fiercely showed a reduction in immunocompetence with respect to control chicks, but the two groups showed no difference in growth rate. The largest and the smallest nestlings in each brood showed a similar response to the treatment. These results strongly suggest a trade-off between begging and immunocompetence in this species. This trade-off may be a consequence either of resources from the immune system being reallocated to begging behaviour, or of adaptive immunosuppression in order to avoid oxidative stress. Steroid hormones are proposed as mediators of such a trade-off.     

Brood size and begging intensity in nestling birds

Theoretical models suggest that sibling competition should selectfor conspicuous begging signals. If so, begging intensity mightbe expected to increase with the number of competitiors. Thepurpose of our study was to examine the relationship betweenbegging intensity and brood size using nestling tree swallows(Tachycineta bicolor) as our model. Over 2 years, we videotapedbegging behavior in unmanipulated broods of different sizes.We found that begging intensity increased with brood size. Theaverage weight of nestlings in each brood did not vary withbrood size, but feeding rate per nestling decreased with broodsize, suggesting that nestlings in larger broods begged moreintensively, possibly because they were hungrier. We also conductedan experiment to examine the effect of nest mates on beggingin different-sized broods. We found that nestlings with similarweights, previous competitive environments, and food deprivationbegged more intensively in large broods than in small broods.Overall, our study indicates that begging intensity increaseswith brood size in tree swallows. This relationship may resultfrom interactions among brood mates rather than from lower feeding rates to individual nestlings in larger broods.