The adaptive value of parental responsiveness to nestling begging

Despite extensive theoretical and empirical research into offspring food solicitation behaviour as a model for parent–offspring conflict and communication, the adaptive value of parental responsiveness to begging has never been tested experimentally. Game theory models, as well as empirical studies, suggest that begging conveys information on offspring state, which implies that parental investment can be better translated to fitness by responding to begging when allocating resources rather than by ignoring it. However, this assumption and its underlying mechanisms have received little or no attention. Here we show by experiments with hand-raised house sparrow (Passer domesticus) nestlings that a ‘responsive parent’ will do better than a hypothetical ‘non-responsive’ mutant (that provides similar food amounts, but irrespective of begging). This is neither because food-deprived nestlings convert food to mass more efficiently, however, nor because responsiveness reduces costly begging. Rather, responsiveness to begging is adaptive because it reduces two opposing risks: one is wasting time when returning too soon to feed already satiated nestlings and the other is repeatedly overlooking some nestlings as a result of the stochastic nature of a random, non-responsive strategy. This study provides the first experimental evidence for the adaptive value of parental responsiveness to offspring begging

Parent–offspring conflict and co-adaptation: behavioural ecology meets quantitative genetics

The evolution of the complex and dynamic behavioural interactions between caring parents and their dependent offspring is a major area of research in behavioural ecology and quantitative genetics. While behavioural ecologists examine the evolution of interactions between parents and offspring in the light of parent–offspring conflict and its resolution, quantitative geneticists explore the evolution of such interactions in the light of parent–offspring co-adaptation due to combined effects of parental and offspring behaviours on fitness. To date, there is little interaction or integration between these two fields. Here, we first review the merits and limitations of each of these two approaches and show that they provide important complementary insights into the evolution of strategies for offspring begging and parental resource provisioning. We then outline how central ideas from behavioural ecology and quantitative genetics can be combined within a framework based on the concept of behavioural reaction norms, which provides a common basis for behavioural ecologists and quantitative geneticists to study the evolution of parent–offspring interactions. Finally, we discuss how the behavioural reaction norm approach can be used to advance our understanding of parent–offspring conflict by combining information about the genetic basis of traits from quantitative genetics with key insights regarding the adaptive function and dynamic nature of parental and offspring behaviours from behavioural ecology

A chemical signal of offspring quality affects maternal care in a social insect

Begging signals of offspring are condition-dependent cues that are usually predicted to display information about the short-term need (i.e. hunger) to which parents respond by allocating more food. However, recent models and experiments have revealed that parents, depending on the species and context, may respond to signals of quality (i.e. offspring reproductive value) rather than need. Despite the critical importance of this distinction for life history and conflict resolution theory, there is still limited knowledge of alternative functions of offspring signals. In this study, we investigated the communication between offspring and caring females of the common earwig, Forficula auricularia, hypothesizing that offspring chemical cues display information about nutritional condition to which females respond in terms of maternal food provisioning. Consistent with the prediction for a signal of quality we found that mothers exposed to chemical cues from well-fed nymphs foraged significantly more and allocated food to more nymphs compared with females exposed to solvent (control) or chemical cues from poorly fed nymphs. Chemical analysis revealed significant differences in the relative quantities of specific cuticular hydrocarbon compounds between treatments. To our knowledge, this study demonstrates for the first time that an offspring chemical signal reflects nutritional quality and influences maternal care

Effect of Variation in Nestling Hunger Levels on the Begging Behaviour of Nestlings and the Provisioning Behaviour of Adult American Kestrels

Little is known about how variation in nestling begging intensity influences the behaviour of adult raptors and how responses of adult males and females to such variation might differ. Our objective was to manipulate the begging intensity of nestling American Kestrels (Falco sparverius) and examine the responses of adults. We studied 12 pairs of American Kestrels nesting in nest boxes from 1 March to 1 July 2014 at the Blue Grass Army Depot, Madison County, Kentucky. Nest boxes were modified with a separate compartment for a camcorder to record nestling behaviour, and a second camcorder was placed outside the nests to monitor adult behaviour. To manipulate nestling hunger levels, 12 to 26-day-old nestlings in six nests were deprived of food for 24 h and those in the other six nests were fed until satiated. At each nest, we alternated control (no treatment) and treatment (fed or food deprived) days over a 4 day period to minimise the possible effect of nestling age on adult and nestling behaviour. Nestling begging intensity differed among treatments, with nestlings in food-deprived nests begging with greater intensity after food deprivation and those in fed-treatment nests begging with less intensity after being fed. Adult male and female American Kestrels provisioned nestlings at similar rates, with both sexes feeding nestlings at higher rates after food deprivation and at lower rates after fed treatments. Thus, the begging behaviour of nestling American Kestrels varied with hunger level, and adult American Kestrels responded by adjusting provisioning rates. Although the response of adults to nestling begging suggests that natural selection might favour \u27dishonest\u27 begging to obtain more food, the potential costs of \u27dishonest\u27 begging, such as attracting predators, reduced immunocompetence, and loss of indirect fitness benefits if such begging negatively impacts siblings and parents, may outweigh any possible benefit

Atmospheric signal propagation

GNSS satellites emit signals which propagate as electromagnetic waves through space to the receivers which are located on or near the Earth’s surface or on other satellites. Thereby, electromagnetic waves travel through the ionosphere and the neutral atmosphere (troposphere) which causes signals to be delayed, damped and refracted as the refractivity index of the propagation media is not equal to one. In this chapter, the nature and effects of GNSS signal propagation in both the troposphere and the ionosphere, is examined. After a brief review of the fundamentals of electromagnetic waves their propagation in refractive media, the effects of the neutral atmosphere are discussed. In addition empirical correction models as well as state-of- the-art atmosphere delay estimation approaches are presented. Effects related to signal propagtion through the ionosphere are dealt in a dedicated section by describing the error contribution of first up to third order terms in the refractive index and ray path bending. After discussing diffraction and scattering phenomena due to ionospheric irregularities, mitigation techniques for different types of applications are presented