鱼类亲代抚育行为的研究进展

亲代抚育行为（parental care behavior）是指动物对其后代或其亲缘后代提供保护和养育的所有活动,属于本能行为的一种,广泛存在于动物界之中。鱼类在其为数不多的科中充分发展了几乎所有类型的亲代抚育行为,因而成为研究该行为的最佳物种之一。随着威廉斯原理（Williams’s Principle）的提出和应用,人们对鱼类亲代抚育行为的探索逐步由定性向定量发展,普遍认同了在鱼类进化中,雄性抚育模式得以占据支配地位的缘由并非是因为雄性在抚育活动中获得了较多的利益,而是由于在获取相同利益时雄性损失的未来投资成本较雌性低的观点。近年来的研究证实,在亲代抚育过程中存在着某种动态调整机制,其中四个比较关键的影响因素分别为：亲本所抚育的子代数量、亲本先前的投资、亲本与被抚育子代间的遗传关联度和亲本未来的交配机会。     

Fishes as models in studies of sexual selection and parental care

Fishes are by far the most diverse group of vertebrates. This fact is in no way, however, reflected in their use as model organisms for understanding sexual selection or parental care. Why is this so? Is it because fishes are actually poor models? The usefulness of fishes as models for sexual selection and parental care is discussed by emphasizing some problems inherent in fish studies, along with a number of reasons why fishes are indeed excellently suited. The pros and cons of fishes as models are discussed mainly by comparison with birds, the most popular model organisms in animal behaviour. Difficulties include a lack of background knowledge for many species, and the problems of marking and observing fishes in their natural environment. Positive attributes include the diversity of lifestyles among fishes, and the ease with which they can be studied experimentally in the laboratory. How useful fish models can be is briefly illustrated by the impressive and broadly relevant advances derived from studies of guppies Poecilia reticulata and three‐spined sticklebacks Gasterosteus aculeatus . A selection of topics is highlighted where fish studies have either advanced or could greatly enhance, the understanding of processes fundamental to animal reproductive dynamics. Such topics include sex role dynamics, the evolution of female ornamentation and mate choice copying. Finally, a number of potential pitfalls in the future use of fish as models for sexual selection and parental care are discussed. Researchers interested in these issues are recommended to make much more extensive use of fish models, but also to adopt a wider range of models among fishes.     

Risk taking during parental care: a test of the harm-to-offspring hypothesis

Amount of risk taking during parental care is often explainedin relation to the reproductive value of the offspring. The"harm-to-offspring hypothesis" focuses on the relative harma period of no parental care can do to the offspring. Accordingto this hypothesis, parents should take greater risks for offspringin poor condition than for offspring in good condition. We manipulatedoffspring condition in the pied flycatcher (Ficedula hypoleuca)and tested the harm-to-offspring hypothesis by exposing parentsto a predator model (a sparrowhawk, Accipiter nisus). Time elapseduntil a parent first entered the nest-box was used as a risk-takingmeasure. Parents spent significantly shorter time until first nestvisit for offspring in poor condition than for offspring ingood condition. Hence, the harm-to-offspring hypothesis wassupported.     

Sexual conflict over parental care promotes the evolution of sex differences in care and the ability to care

Strong asymmetries in parental care, with one sex providing more care than the other, are widespread across the animal kingdom. At present, two factors are thought to ultimately cause sex differences in care: certainty of parentage and sexual selection. By contrast, we here show that the coevolution of care and the ability to care can result in strong asymmetries in both the ability to care and the level of care, even in the absence of these factors. While the coevolution of care and the ability to care does not predict which sex evolves to care more than the other, once other factors give rise to even the slightest differences in the cost and benefits of care between the sexes (e.g. differences in certainty in parentage), a clear directionality emerges; the sex with the lower cost or higher benefit of care evolves both to be more able to care and to provide much higher levels of care than the other sex. Our findings suggest that the coevolution of levels of care and the ability to care may be a key factor underlying the evolution of sex differences in care.     

Life history evolution in cichlids 1: revisiting the evolution of life histories in relation to parental care

Empirical links between egg size and duration of parental care in fishes have generated a considerable amount of theory concerning life history evolution. However, to date, this link has not been investigated in relation to other important life-history traits such as clutch size and body size, or while controlling for shared ancestry between species. We provide the first phylogenetically based tests using a database with information on egg size, clutch size, body size and care duration in cichlid fishes (Cichlidae). Multiple regression analyses, based on independent contrasts on both the species and the genus level, showed that clutch size is the variable most closely related to duration of care. This pattern appeared to be driven by post-hatch care relationships. Our results show that, contrary to expectation, there is no positive link between egg size and care duration in Cichlidae. Instead, greater reproductive output through increased clutch size investment appears to have coevolved with greater care of offspring. We suggest that re-evaluation of the generality of current models of the evolution of egg size under parental care in fishes is needed.     

Primitive bony fishes, with especial reference to Cheirolepis and palaeonisciform actinopterygians

The relationships of the Devonian palaeonisciform fish Cheirolepis are examined and the early evolutionary trends within the Actinopterygii and the Osteichthyes are considered.
Cheirolepis is the most primitive known actinopterygian. The contemporary stegotrachelid palaeonisciforms are more advanced in their cranial and locomotor anatomy. The general directions of these advances are similar to those subsequently displayed by later palaeonisciforms over the stegotrachelids themselves. Cheirolepis , furthermore, possesses many characters which can be logically interpreted as primitive for the Osteichthyes by extrapolation of trends in actinopterygian and sarcopterygian lineages. 11 is the most primitive known osteichthyan.
The Osteichthyes are considered to have arisen from a micromerically-scaled acanthodian or acanthodian-like ancestor at the end of the Silurian period.     

Seasonal variation in parental care, offspring development, and reproductive success in the burying beetle, Nicrophorus vespillo

1. Beetles of the genus Nicrophorus reproduce on small vertebrate carcasses that they bury in the soil to provide the larvae with food. Usually, both parents cooperate in brood care by feeding and guarding their progeny. 2. In pairs of the common European species N. vespillo, the duration of care depended on the time of year when the beetles reproduced. Both in 1990 and in 1991, male and female parents stayed longer with their broods when reproduction started in spring than when reproduction started in early or late summer. This was probably due to the longer development time of the larvae caused by lower temperatures in spring, because laboratory experiments suggested a strong influence of temperature on both the duration of brood care and offspring development. 3. The number of adult offspring produced by a beetle pair did not vary among different times of the year. 4. The median time required for offspring development, measured as time from burial of the carcass to emergence of young adults, was between 62 and 84 days. When the beetles reproduced in late summer, only about three-quarters of the offspring left the soil and hibernated as adults. The remaining offspring stayed underground and adults appeared on the soil surface the following spring. They still showed the flexible cuticle typical of newly-hatched beetles, suggesting that they may have overwintered in a pre-adult stage.     

The general protected invasion theory: Sex biases in parental and alloparental care

The biases towards eusociality, female workers and maternal care in haplodiploid versus diploid insects may result from the relatively low probabilities that rare mutant, partially dominant alleles promoting these behaviours will be lost by genetic drift in haplodiploid populations (Reeve, 1993). A generalization of this 'protected invasion' theory also predicts that parental and alloparental care will tend to be associated with the homogametic sex in diploid populations if the Y chromosome of the heterogametic sex is absent or largely inert. Sex differences in (allo)parental care (i.e. either parental or alloparental care) should increase with increased asymmetry between the sexes in the fraction of behaviour-influencing loci occurring on their characteristic sex chromosomes. The theory explains the strong predisposition towards female (allo)parental care in mammals, a contrasting tendency towards male (allo)parental care in birds, the propensity for joint male and female (allo)parental care in termites, and biases towards female cooperation in social spiders. The theory also explains the apparent rarity or absence of alloparental care in marsupials, an intriguing consequence of preferential paternal X-chromosome inactivation in this taxon. Thus protected invasion theory possibly provides new insights into the relationship between social structure and the genetic system. The theory does not compete with ecological or kin-selective hypotheses for the advantages of (allo)parental care; indeed, such advantages must exist for protected-invasion biases to operate.     

Subsocial behaviour in Oniticellus cinctus (Coleoptera, Scarabaeidae): effect of the brood on parental care and oviposition

ABSTRACT. In Oniticellus cinctus (F.) the nest chambers each contain about twenty brood balls. Females enlarge the brood balls during the egg and larval stages and remain in the chamber for the whole period of brood development (1 month); they then make a new nest after 1 week. The presence of the brood releases parental care and ensures that the mother remains in the nest: she repairs defects in the brood balls and the nest, and expels other O. cinctus females. A new ball is formed around a naked O. cinctus larva, but larvae of other species are killed. In addition, the brood inhibits oviposition: removal (or addition) of brood balls stimulates (or inhibits) egg laying. In inhibited ovarioles, existing follicles are resorbed and production of new ones ceases. Control of clutch size by the brood is an adaptation to the nest structure and life history of O. cinctus. It may have an important role in the reproductive strategy of other insects with parental care.     

Copulation, spawning and parental care in captive ocean pout

The ocean pout copulates through direct genital contact for internal fertilization. A complete spawning event consists of copulation, oviposition, and the female displaying parental care by wiping and wrapping herself around the eggs.     

Energetics of parental care in six syntopic centrarchid fishes

We studied parental behavior in six syntopically breeding species of centrarchid fishes to determine whether energetic costs could contribute to our understanding of the diversity of parental care. We used a combination of underwater videography, radio telemetry and direct observation to examine how the cost of parental care varied with both its duration and intensity. Duration of parental care, activity patterns, and energetic costs varied widely among species. Overall, the duration of care increased with parental size between species. When energetic costs were adjusted for species-specific differences in the duration of parental care, the cost of parental care also increased with mean size of the species. Species with extended parental care exhibited stage-specific patterns of activity and energy expenditure consistent with parental investment theory, whereas fish with short duration parental care tended to maintain high levels of activity throughout the entire period of parental care. The only apparent exception (a species with brief parental care but stage-specific behavior) was a species with multiple breeding bouts, and thus effectively having protracted parental care. These data suggest that some species with short duration parental care can afford not to adjust parental investment over stages of offspring development. Using our empirical data on parental care duration and costs, we reevaluated the relationship between egg size and quality of parental care. Variation in egg size explained almost all of the observed variation in total energetic cost of parental care, and to a lesser degree, duration—the larger the eggs, the more costly the parental care. This research highlights the value of incorporating energetic information into the study of parental care behavior and testing of ecological theory.     

Sexual conflict over parental investment in repeated bouts: negotiation reduces overall care

Understanding the evolution of parental care is complicated by the occurrence of evolutionary conflicts of interest within the family, variation in the quality and state of family members, and repeated bouts of investment in a family of offspring. As a result, family members are expected to negotiate over care. We present a model for the resolution of sexual conflict in which parents negotiate over repeated bouts of care. Negotiation is mediated by parents deciding at the start of each bout how much care to give on the basis of the state (mass) of offspring, which reflects the amount of care previously received. The evolutionarily stable pattern of care depends on whether the parents care together for the whole family, or each cares alone for part of the divided family. When they care together, they provide less care in the first bout, more in the last bout, and less care overall, resulting in lower parental and offspring fitness. Our results emphasize that negotiation over parental care may occur as a means of avoiding exploitation owing to sexual conflict, even in the absence of variation in the quality of either sex of parent, and lead to a reduction in fitness.     

The evolution of human biosocial behaviour

The analysis of human behaviour can be approached from three different but complementary perspectives: the first includes the eternal debate about the degree of environmental or genetic determinism of social behaviour; the second, in the context of evolutionary ecology, concerns the evaluation of behavioural responses to morphological and/or environmental changes that have been the key to success in our species; the third, in the context of the analysis of the biology of the health of modern day populations, deals with the biological consequences of social behavioural changes. The secret of the success of a species resides in its capacity to respond behaviourally to the morphological and environmental changes produced during its biological history. What is unique about humans concerns our brains and their derived function: flexible behaviour. The morphological, physiological and behavioural changes that allow thecreation and maintenance of such a large brain are intimately connected to reproduction and therefore to the biosociology of women, the members of the speciesHomo sapiens in whom are combined, with notable success, a prolonged life-cycle, some anatomical and physiological traits and flexible reproductive behaviour, which together confer a decisive demographic advantage on them over other hominids. In summary, as well as contributing to the spread of this information by which women can make informed decisions based on a knowledge of causes, with the aim of optimizing our health it is necessary to consider the possibility of approaching some aspects of our biology and behaviour as patterns fixed by evolution. Three factors would be the most easily adjustable for this purpose: a) to delay the age of sexual maturation, b) to bring forward the age of first maternity, and c) to encourage breastfeeding on demand without substitutes and for a 3–5 month period.     

Quantifying the costs and benefits of parental care in female treehoppers

Parental protection of eggs represents one of the most basicforms of parental care. Theory suggests that even such basicparental investment represents a trade-off between current offspringsurvival and future reproductive success. However, few studieshave quantified the underlying costs and benefits of parentalcare for marked individuals across an entire lifetime. I markedand followed 370 females of Publilia concava (Hemiptera: Membracidae)that exhibited a range of guarding durations for their firstclutch. Greater hatching success was correlated with longerguarding durations, and a removal experiment verified that femalepresence was responsible for a twofold increase in hatchingsuccess. On the other hand, females that remained to guard eggshad a lower number and size of future broods, suggesting thatparental care may reduce lifetime fecundity. Marked femalesexhibited a bimodal distribution of guarding durations, reflectingthe extreme tactics of immediate abandonment or remaining throughhatching. Estimates of lifetime number of nymphs produced byfemales that abandon eggs early versus guard eggs through hatchingrevealed roughly equivalent levels of fitness. I discuss theconditions under which we might expect a female to adopt eachof the alternative tactics, given the costs and benefits ofparental care that were quantified in this study.     

Extended parental care and delayed dispersal: northern, tropical, and southern passerines compared

Using modern comparative methods, we found that both time toindependence and time with parents were significantly longerin southern hemisphere and tropical birds than in northern hemisphereones. These differences held even after removing Australianpasserines or cooperatively breeding species, and they do notdepend on habitat, diet, or migration pattern. In southern hemisphereand tropical regions, both cooperative breeding and non-cooperativeparents continue to feed their young for a similar length oftime, but cooperative breeders allow them to stay longer intheir natal territory after they become nutritionally independent.Nevertheless, the young of non-cooperative species stay longerwith their parents than do the young of non-cooperative speciesin the temperate northern hemisphere. The fact that extendedperiods of post-fledging parental care are widespread amongpasserines provides further empirical support for the view thatlife histories of southern and tropical birds are ‘slow,’with small clutches, extended parental care, and long lifespan;parents take care of fewer young for longer. These results supportrecent theoretical models that predict that high adult survivaland low turnover of territory owners generally favor natal philopatry.We suggest that the reasons why some species (with or withoutcooperative breeding) exhibit natal philopatry and others donot lie in the balance between productivity and survival ofadults and of retained or dispersing offspring.     

Brood desertion in Kentish plover: the value of parental care

To understand the evolution of parental care, one needs to estimatethe payoffs from providing care for the offspring and from terminatingcare and deserting them. In this study we estimated the payofffrom care provision, and in a companion paper we analyze thepayoff from offspring desertion. In the current study we experimentallyinvestigated the influence of the number and sex of attendingparents on growth and survival of offspring in the Kentish ploverCharadrius alexandrinus, in two sites (A and B). Either the maleor the female parent was removed from some broods at hatchingof the chicks (female-only and male-only broods, respectively),whereas in control broods both parents were allowed to attendtheir young. At site A survival of the chicks was lower in uniparental(male-only and female-only) broods than in control broods, whereaswe found no difference in brood survival at site B. Brood survivaldecreased over the season. Removal of either parent did not influencethe growth of the young, although growth varied over the breeding season,and it was significantly different between the sites. Theseresults suggest that the payoff from parental care decreasesover the breeding season and that the value of parental care(i.e., the contribution of parents to the survival of theiryoung) may depend on the environment.     

Parental behaviour in Copris lunaris (Coleoptera, Scarabaeidae): care and defence of brood balls and nest

Abstract. 1. Nesting female beetles righted brood balls (so as to replace the egg or larva in the uppermost position) and repaired damaged balls. This behaviour required the presence of an egg or larva in the ball, or of a short-lasting material found just after oviposition. The shape of the ball was also a righting stimulus since artificial ellipsoids were stood on end.
2. Balls containing dichloromethane extracts of C.lunaris brood were righted and repaired. Eggs and larvae of several other Scarabaeidae did not release these responses but were destroyed.
3. Righting behaviour was released when brood was absent from the top of the ball. The beetle then crawled vertically downwards and, if it encountered the displaced apex, a novel rolling action followed which automatically turned the ball towards the correct position.
4. An opening made in the nest was repaired with soil excavated from the chamber floor. Clunaris adults and Aphodius fossor larvae were attacked if they were encountered in the nest.     

Effects of among-offspring relatedness on the origins and evolution of parental care and filial cannibalism

Parental care is expected to increase the likelihood of offspring survival at the cost of investment in future reproductive success. However, alternative parental behaviours, such as filial cannibalism, can decrease current reproductive success and consequently individual fitness. We evaluate the role of among-offspring relatedness on the evolution of parental care and filial cannibalism. Building on our previous work, we show how the evolution of care is influenced by the effect of among-offspring relatedness on both the strength of competition and filial cannibalism. When there is a positive relationship between among-offspring competition and relatedness, parental care will be favoured when among-offspring relatedness is relatively low, and the maintenance of both care and no-care strategies is expected. If the relationship between among-offspring competition and relatedness is negative, parental care is most strongly favoured when broods contain highly related offspring. Further, we highlight the range of conditions over which the level of this among-offspring relatedness can affect the co-occurrence of different care/no care and cannibalism/no cannibalism strategies. Coexistence of multiple strategies is independent of the effects of among-offspring relatedness on cannibalism but more likely when among-offspring relatedness and competition are positively associated.