On the oxidative cost of begging: antioxidants enhance vocalizations in gull chicks

Noguera, J.C., Morales, J., Pérez, C. & Velando, A., 2010. On the oxidative cost of begging: antioxidants enhance vocalizations in gull chicks. Behavioral Ecology, 21(3), pp.479-484.

Offspring solicit food to their parents by begging displays, which are important in the parent–offspring communication. Most theoretical approximations on this behavior have centered on the view of begging as an honest signal of need or as a form of scramble competition for resources. In both signaling models, costly begging is necessary to stabilize the begging strategy at equilibrium. Nevertheless, evidence supporting begging as costly behavior remains scarce. We investigated whether oxidative stress may represent a general form of proximate cost of begging and also whether begging is related to offspring nutritional condition. To test this, we experimentally modified the chicks’ nutritional condition and vitamin E availability and measured the effects on different begging components. The intensity of all begging components increased in chicks that were intake restricted, whereas vitamin E specifically enhance the total number of chatter calls given by chicks, mainly in those with a lower body size. Our results suggest that begging behavior is an antioxidant demanding activity and support the idea that oxidative stress may be a cost of begging. Our findings also suggest that begging behavior may be an honest signal of the nutritional and oxidative status of the chicks.

The evolution of multicomponent begging display in gull chicks: sibling competition and genetic variability

Kim, S.Y., Noguera, J.C., Morales, J. & Velando, A., 2011. The evolution of multicomponent begging display in gull chicks: sibling competition and genetic variability. Animal Behaviour, 82(1), pp.113-118.

The evolution of begging display may be influenced by gene–environment interaction, through the mechanisms that adjust begging behaviour to environmental conditions of offspring, including intensity of sibling competition within broods. We decomposed the complex begging display of yellow-legged gull, Larus michahellis, chicks into two different functional components: begging for food (pecks) and drawing the attention of parents (chatter calls). We examined these begging components in 2-day-old chicks that hatched and grew up in foster nests, by performing a begging test for each chick alone without the hindrance of its foster siblings. Male chicks and those with poorer body condition begged for food at higher rates than females and those with better body condition, respectively. Chicks from larger broods begged for food more frequently, but chicks from male-biased broods begged less frequently. If begging is costly, chicks may adjust their begging efforts to the intensity of sibling competition. Frequency of chatter calls varied with sex, chick order within broods and body condition: females, the third chicks and those with poorer condition produced chatter calls more frequently. Genetic origin had a significant effect on frequency of chatter calls but not on begging for food, while foster nest effect was null in both traits. Therefore, chatter calls (but not pecks) can be subject to evolution under directional selection. Different begging components may have evolved through different evolutionary pathways.

Hatching hierarchy but not egg-related effects governs behavioral phenotypes in gull chicks

Diaz-Real, J., Kim, S.Y. & Velando, A., 2016. Hatching hierarchy but not egg-related effects governs behavioral phenotypes in gull chicks. Behavioral Ecology. doi: 10.1093/beheco/arw110

In many bird species that practice parental care, siblings often compete for resources and care provided by their parents, although their strategies differ according to hatching rank and condition. Differences in offspring strategies are generally attributed to hatching order and maternal effects, which are difficult to separate because these effects are often correlated. For example, third-hatched chicks of large gull species receive more egg testosterones and corticosterone, which influence early behavioral patterns. In this study, we carried out a cross-fostering experiment with first- and last-laid eggs of the yellow-legged gull (Larus michahellis) to test whether the within-brood variation in behavioral strategies for competing with siblings and coping with stress are due to maternal effects or to hatching order. Chicks hatched in the last position within the experimental brood emitted more chatter calls to attract parents’ attention, were less prone to respond to warning of danger, and had a lower breathing rate while restrained than first-hatched chicks. Egg laying order did not affect chick behaviors or breathing rate. Thus, we concluded that the different behavioral strategies of chicks were determined by their posthatching experience and not by the original egg position within the clutch. Last-laid eggs were smaller and chicks from those eggs grew slower than chicks from first-laid eggs. Independently of the original laying order, chicks that hatched first in the experimental brood grew faster than their siblings. Overall, our results indicate that behavioral strategies of chicks are plastic and influenced by their early social

Begging response of gull chicks to the red spot on the parental bill

Velando, A., Kim, S.-Y. & Noguera, J. C. Begging response of gull chicks to the red spot on the parental bill. Animal Behaviour 85, 1359–1366 (2013).

In some animals, offspring begging is elicited by parents through behavioural or morphological signals. The red spot on the lower mandible in adult gulls is one of the best-known examples of a signal triggering chick begging. We examined whether the begging response of chicks (pecking for food and the chatter call for drawing parental attention) was affected by the spot size within the natural range of variation on a dummy head. Using a cross-fostering experiment, we examined whether these responses covary with the size of the genetic or social parent’s spot. We found that the natural variation in size of this parental signal strongly influenced intensity of chick begging. Pecking increased when chicks were stimulated by a larger red spot. Additionally, pecking intensity increased in chicks reared by mothers with a large red spot, suggesting that this begging component is influenced by previous experience. In contrast, chick hatching order affected the number of chatter calls produced in relation to the size of the red spot on the dummy, suggesting the presence of different begging strategies according to brood hierarchy. The differential call response to a small/large red spot on the dummy was positively correlated with the original mothers’ red spot size and negatively with that of the original fathers. These results suggest a genetic correlation between biased chick response for a large spot and parental signal in contrasting patterns for mothers and fathers. Our results suggest that the parental red spot and offspring begging are traits subject to coevolution.

Maternal testosterone influences a begging component that makes fathers work harder in chick provisioning

Noguera, J. C., Kim, S.-Y. & Velando, A. Maternal testosterone influences a begging component that makes fathers work harder in chick provisioning. Hormones and behavior 64, 19–25 (2013).

In species with biparental care, parents disagree evolutionarily over the amount of care that each of them is willing to provide to offspring. It has recently been hypothesised that females may try to manipulate their mates by modifying offspring begging behaviour through yolk hormone deposition, shifting the division of labour in their own favour. To test this hypothesis we first investigated how yellow-legged gull (Larus michahellis) parents feed offspring in relation to each component of complex begging behaviour and if feeding behaviour varies between sexes. Then we investigated the effect of yolk testosterone on chicks’ begging by experimentally increasing yolk testosterone levels. Our results revealed that yolk testosterone has a component-specific effect on chicks’ begging, specifically increasing the number of chatter calls. Parental feeding effort was influenced by the number of chatter calls emitted by chicks, but most importantly, the influence was stronger in male than in female parents. Moreover, chick body mass increased with the number of paternal feeds. In conclusion, these results show that female gulls may use yolk testosterone deposition to exploit their partners as predicted by the ‘Manipulating Androgen Hypothesis (MAH)’.