What are the benefits of parental care? The importance of parental effects on developmental rate

The evolution of parental care is beneficial if it facilitates offspring performance traits that are ultimately tied to offspring fitness. While this may seem self‐evident, the benefits of parental care have received relatively little theoretical exploration. Here, we develop a theoretical model that elucidates how parental care can affect offspring performance and which aspects of offspring performance (e.g., survival, development) are likely to be influenced by care. We begin by summarizing four general types of parental care benefits. Care can be beneficial if parents (1) increase offspring survival during the stage in which parents and offspring are associated, (2) improve offspring quality in a way that leads to increased offspring survival and/or reproduction in the future when parents are no longer associated with offspring, and/or (3) directly increase offspring reproductive success when parents and offspring remain associated into adulthood. We additionally suggest that parental control over offspring developmental rate might represent a substantial, yet underappreciated, benefit of care. We hypothesize that parents adjust the amount of time offspring spend in life‐history stages in response to expected offspring mortality, which in turn might increase overall offspring survival, and ultimately, fitness of parents and offspring. Using a theoretical evolutionary framework, we show that parental control over offspring developmental rate can represent a significant, or even the sole, benefit of care. Considering this benefit influences our general understanding of the evolution of care, as parental control over offspring developmental rate can increase the range of life‐history conditions (e.g., egg and juvenile mortalities) under which care can evolve.     

Should advertising parental care be honest?

Species with paternal care show less exaggerated sexual ornamentation than those in which males do not care, although direct benefits from paternal care can vastly exceed the indirect benefits of mate choice. Whether condition-dependent handicaps can signal parenting ability is controversial. The good-parent process predicts the evolution of honest signals of parental investment, whereas the differential-allocation model suggests a trade-off between the attractiveness of a mate and his care-provisioning. I show that both alternatives can arise from optimal allocations to advertisement, parental investment and future reproductive value of the male, and that the male''s marginal fitness gain from multiple matings determines which option should apply. The marginal gain is diminishing if opportunities for polygyny or extra-pair copulations are limited. Advertisement is then expected to be modest and honest, indicating genetic quality and condition-dependent parental investment simultaneously. Increasing marginal gains are likely to be related to cases where genetic quality has a significant influence on offspring fitness. This alternative leads to differential allocation with stronger advertisement, more frequent extra-pair copulations, and diminished male care. Reliability is also reduced if allocation benefits have thresholds, e.g. if there is a minimum body condition required for survival, or if females use a polygyny-threshold strategy of mate choice.     

Evolution of eusocial behavior: Offspring choice or parental parasitism?

Assuming the subsocial route towards eusociality, I show that selection favors worker habits (under haplodiploidy) provided the mutant workers are able to bias the sex ratio towards sisters and/or selectively substitute their sons for brothers. If the original workers are not able to do this, selection does not favor the habit. However, under these conditions, selection is indifferent as to whether an individual rears offspring or sibs. This makes it easy for a mother to enlist her daughter's services in rearing other offspring, since the daughter cannot evolve to stop the parental parisitism. These results deal with the origin of eusocial behavior. I also look at selection acting on genes (to invest in offspring rather than sibs) in existing eusocial societies. It is shown that selection for laying workers is very strong, even if such workers give up rearing a seemingly advantageous combination of brothers and sisters. This poses distinct problems for the maintenance of eusocial societies.     

Phenotypic plasticity, parental effects, and parental care in plants? I. An examination of spike reflectance in Plantago lanceolata (Plantaginaceae)

We explore the relationships among phenotypic plasticity, parental effects, and parental care in plants by presenting data from four experiments examining reflectance/color patterns in Plantago lanceolata. In three experiments, we measured spike (inflorescence) reflectance between 362 and 850 nm using a spectrophotometer with an integrating sphere. Experiments show that (1) spike reflectance changes seasonally within and outside the visible portion of the spectrum of radiant energy, (2) increasing ambient temperature causes an individual plant to produce flowering and fruiting spikes that reflect more/lighten in color (the greatest changes occur in the regions around 550 nm and between 750 and 850 nm, the visible and near-infrared regions, respectively), (3) responses are reversible, (4) genotypes within populations and populations from different latitudes differ in mean reflectance and degree of phenotypic plasticity. In a fourth experiment, we measured internal spike temperature. Darker spikes, those produced at lower temperature, got hotter than did lighter spikes in full sun. Thus, plants can partially thermoregulate reproduction and the embryonic development of their offspring. In light of a previous experiment, data suggest that thermoregulation produces adaptive parental effects and is a mechanism by which P. lanceolata provides parental care.     

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.     

How do changes in parental investment influence development in echinoid echinoderms?

SUMMARY Understanding the relationship between egg size, development time, and juvenile size is critical to explaining patterns of life-history evolution in marine invertebrates. Currently there is conflicting information about the effects of changes in egg size on the life histories of echinoid echinoderms. We sought to resolve this conflict by manipulating egg size and food level during the development of two planktotrophic echinoid echinoderms: the green sea urchin, Strongylocentrotus droebachiensis and the sand dollar, Echinarachnius parma . Based on comparative datasets, we predicted that decreasing food availability and egg size would increase development time and reduce juvenile size. To test our prediction, blastomere separations were performed in both species at the two-cell stage to reduce egg volume by 50%, producing whole- and half-size larvae that were reared to metamorphosis under high or low food levels. Upon settlement, age at metamorphosis, juvenile size, spine number, and spine length were measured. As predicted, reducing egg size and food availability significantly increased age at metamorphosis and reduced juvenile quality. Along with previous egg size manipulations in other echinoids, this study suggests that the relationship between egg size, development time, and juvenile size is strongly dependent upon the initial size of the egg.     

Are stepchildren over-represented as victims of lethal parental violence in Sweden?

Evolutionary psychologists have suggested that stepchildren should be over-represented as victims of lethal parental violence compared with children living with their two genetic parents, because of relatively more lapses in parental solicitude among stepparents. In our study, using data over a period of 35 years in Sweden (1965-1999), there was no overall over-representation of stepchildren as victims. For very young stepchildren there was a tendency for over-representation. In families with both stepchildren and children genetically related to the offender, genetic children tended to be more likely to be victims.     

The cost of parental care inHaplochromis ‘argens’ (Cichlidae)

Synopsis The cost of parental care in the maternal, mouthbrooding cichlid,Haplochromis ‘argens’ was measured in terms of parental survival, fecundity and breeding frequency. In comparison with non-parental females, those that mouthbrooded eggs for a period of 16 days took 33 % longer to respawn. Parental females also grew more slowly between spawnings, and this may affect their future fecundity. These costs were attributed to the cessation of feeding by parental females while mouthbrooding and/or the hormonal inhibition of oogenesis.     

Do the different parental 'heteromes' cause genomic shock in newly formed allopolyploids?

Allopolyploidy, the joining of two parental genomes in a polyploid organism with diploid meiosis, is an important mechanism of reticulate evolution. While many successful long-established allopolyploids are known, those formed recently undergo an instability phase whose basis is now being characterized. We describe observations made with the Arabidopsis system that include phenotypic instability, gene silencing and activation, and methylation changes. We present a model based on the epigenetic destabilization of genomic repeats, which in the parents are heterochromatinized and suppressed. We hypothesize that loss of epigenetic suppression of these sequences, here defined as the heterome, results in genomic instability including silencing of single-copy genes.     

Components of care: are they honest signals of parental care quality?

It has been argued that male parental care provides direct benefits to females and therefore should be under sexual selection. Given this, we expect signals that honestly indicate the quality of care to be favoured by selection. One such potential signal is care itself. Fish have several features that make them excellent model systems for studying the evolution and dynamics of parental care. We use the flagfish, Jordanella floridae , as a model to evaluate these general ideas. Males of this species guard, clean and fan empty nests and then eggs. Females prefer males that fan more (1) before spawning and (2) when eggs are newly received. When single males and females were paired, males that fanned and visited their nests more prior to spawning were more likely to be mated. Furthermore, among successful males, rates of fanning in the first day after spawning were correlated with the number of eggs received in the future (but not current egg numbers). We then considered whether these two putative signals were correlated and whether males that fan more in these contexts actually have higher egg survivorship. We found no correlation between nest fanning rates before and after spawning and neither 'signal' was predictive of variation in egg survivorship among mated males. We further considered whether pre‐spawning fanning rates were predictive of hatching success in an experiment in which single males were allowed to establish nests and provided eggs. We found little evidence that fanning is an honest signal of care quality and discuss alternative explanations. In particular, we discuss patterns of care elaboration in light of our results.     

Food provisioning to nestling shearwaters: why parental behaviour should be monitored?

We examined temporal variation in food delivery to nestling Cory's shearwaters Calonectris diomedea, by repeated periodic weighings during the night. We tested whether the magnitude and frequency of meals were influenced by the condition of chicks. In contrast to previous studies of chick provisioning in petrels and shearwaters, the evidence of feeding derived from chick weight gains was complemented by data provided by an electronic system, which logged the entry of each parent to the nest. Estimates of feed size and visiting frequency obtained from chick weighing alone differed from similar estimates obtained using the automatic logging equipment. The data obtained with the logging system combined with chick weighing also showed that, to some extent, food provisioning was regulated, chicks left in poorer condition being more likely to receive food the next night than those left in better condition. The methods based on chick weight gains alone did not detect this regulation effect. Our findings suggest that resolving parental visits to the nest is crucial to obtain accurate parameter estimates, and to address the problem of regulation of provisioning rates in Procellariiformes. Our results do not support the hypothesis that accumulation of fat is just a by-product of chronic overfeeding arising from stochastic variation in foraging success at sea. Copyright 1999 The Association for the Study of Animal Behaviour.     

How does parental role influence the activity and movements of breeding wolves?

We studied the activity and movements of a male and a female wolf during breeding in the Bieszczady Mountains, Poland. The female was less active and mobile and on average closer to the den than the male. The male was less active and mobile when the pack size was seven compared to a year when the pack consisted only of the breeding pair. We conclude that the roles played by breeding males and females rearing pups influence their activity, and that breeding males in larger packs move less during the nursing period because of help by other pack members.     

Does genetic background moderate the association between parental education and school achievement?

This study was conducted with a purpose to examine whether the T102C polymorphism of the serotonin receptor 2A (HTR2A) gene moderates the association between parental education and children's school achievement across nine compulsory school years. The study was carried out in a population‐based sample of Finnish students (aged 9, 12 and 15 years, n = 982). It was found that the HTR2A gene was not related to the school achievement at any school level, but moderated the association between maternal education and the children's grade point averages. The T/T genotype carriers benefited most from high‐maternal education, and suffered from a low one more than the carriers of the other variants of the HTR2A gene. The present finding may at least partly answer the important question why academic outcomes of environmental interventions vary even at the same intelligence levels of the students.     

Quantity and quality of communication during parental deployment: Links to adolescents’ functioning

Using an online survey methodology, we examined individual differences in distance communication between 75 adolescents and their deployed parents and found substantial individual differences in both the quantity and quality of their communication. We also examined the statistical associations between these features of distance communication and adolescents’ functioning, including emotional reactions following communication, health-related quality of life, and externalizing and internalizing problems. The quantity of communication of deployed parents with their adolescents was not associated with adolescents’ functioning, but more positive and less controlling communication was statistically associated with adolescents’ higher functioning. Implications for theory, practice, and future research are discussed.     

Visible and “cryptic” segregation of parental chromosomes in embryonic stem hybrid cells

Chromosome segregation of the parental chromosomes was studied in 20 interspecific hybrid clones obtained by fusion of Mus musculus embryonic stem cells with Mus caroli splenocytes. FISH analysis with labeled species specific probes and microsatellite markers was used for identification of the parental chromosomes. Cytogenetic analysis has shown significant intra- and interclonal variability in chromosome numbers and ratios of the parental chromosomes in the hybrid cells: six clones contained all M. caroli chromosomes, nine clones showed moderate segregation of M. caroli chromosomes (from 1 to 7), and five clones showed extensive loss of M. caroli chromosomes (from 12 to complete loss of all M. caroli autosomes). Both methods demonstrated cryptic segregation of the somatic partner chromosomes. For instance, five clones with near-tetraploid chromosome sets contained only few M. caroli chromosomes (from 1 to 8). The data obtained suggest that the tetraploid chromosome set per se is not a sufficient criterion for conclusion on the absence of chromosome loss in the hybrid cells. Note that cryptic chromosome segregation occurred at a high frequency in the examined hybrid clones. Thus, cryptic segregation should be borne in mind for assessing pluripotency and genome reprogramming of embryonic stem hybrid cells.__________Translated from Ontogenez, Vol. 36, No. 2, 2005, pp. 151–158.Original Russian Text Copyright © 2005 by Pristyazhnyuk, Temirova, Menzorov, Kruglova, Matveeva, Serov.     

Are you my mother? Bayesian phylogenetic inference of recombination among putative parental strains

Reconstructing evolutionary relationships using Bayesian inference has become increasingly popular due to the ability of Bayesian inference to handle complex models of evolution. In this review we concentrate on inference of recombination events between strains of viruses when these events are sporadic, ie rare relative to point mutations. Bayesian inference is especially attractive in the detection of recombination events because it allows for simultaneous inferences about the presence, number and location of crossover points and the identification of parental sequences. Current frequentist recombination identification falls into a sequential testing trap. The most likely parental sequences and crossover points are identified using the data and then the certainty of recombination is assessed conditional on this identification. After briefly outlining basic phylogenetic models, Bayesian inference and Markov chain Monte Carlo (MCMC) computation, we summarise three different approaches to recombination detection and discuss current challenges in applying Bayesian phylogenetic inference of recombination.     

How is sexual conflict over parental care resolved? A meta‐analysis

Biparental care of offspring is both a form of cooperation and a source of conflict. Parents face a trade‐off between current and future reproduction: caring less for the current brood allows individuals to maintain energy reserves and increase their chances of remating. How can selection maintain biparental care, given this temptation to defect? The answer lies in how parents respond to changes in each other’s effort. Game‐theoretical models predict that biparental care is evolutionarily stable when reduced care by one parent leads its partner to increase care, but not so much that it completely compensates for the lost input. Experiments designed to reveal responses to reduced partner effort have mainly focused on birds. We present a meta‐analysis of 54 such studies, and conclude that the mean response was indeed partial compensation. Males and females responded differently and this was in part mediated by the type of manipulation used.