Among‐individual heterogeneity in maternal behaviour and physiology affects reproductive allocation and offspring life‐history traits in the garter snake Thamnophis elegans

Accumulating evidence suggests that within‐individual plasticity of behavioural and physiological traits is limited, resulting in stable among‐individual differences in these aspects of the phenotype. Furthermore, these traits often covary within individuals, resulting in a continuum of correlated phenotypic variation among individuals within populations and species. This heterogeneity, in turn, affects individual fitness and can have cross‐generational effects. Patterns of trait covariation, among‐individual differences, and subsequent fitness consequences have long been recognized in reptiles. Here, we provide a test of patterns of among‐individual heterogeneity in behaviour and physiology and subsequent effects on reproduction and offspring fitness in the garter snake Thamnophis elegans. We find that measures of activity levels vary among individuals and are consistent within individuals in reproductive female snakes, indicating stable behavioural phenotypes. Blood hormone and glucose concentrations are not as stable within individuals, indicating that these traits do not describe consistent physiological phenotypes. Nonetheless, the major axes of variation in maternal traits describe behavioural and physiological phenotypes that interact to predict offspring body condition and mass at birth. This differential allocation of energy to offspring, in turn, strongly influences subsequent offspring growth and survival. This pattern suggests the potential for strong selection on phenotypes defined by behaviour–physiology interactions.     

Feral cattle of Swona, Orkney Islands

The feral cattle, currently numbering 33, which inhabit the Orkney island of Swona, are described. Their behaviour while foraging for seaweed is compared with that of sheep, and aspects of their social behaviour and population structure are compared with what has been observed among the Chillingham white cattle.     

Genome-scale analysis of demographic history and adaptive selection

One of the main topics in population genetics is identification of adaptive selection among populations. For this purpose, population history should be correctly inferred to evaluate the effect of random drift and exclude it in selection identification. With the rapid progress in genomics in the past decade, vast genomescale variations are available for population genetic analysis, which however requires more sophisticated models to infer species’ demographic history and robust methods to detect local adaptation. Here we aim to review what have been achieved in the fields of demographic modeling and selection detection. We summarize their rationales, implementations, and some classical applications. We also propose that some widely-used methods can be improved in both theoretical and practical aspects in near future.     

Population and behaviour genetics of Drosophila ananassae

Drosophila ananassae is a cosmopolitan and domestic species. It occupies a nuique status among the Drosophila species due to certain peculiarities in its genetic behaviour. The most unusual feature of this species is spontaneous male recombination in appreciable frequency. The present review summarises the work done on population and behaviour genetics of D. ananassae from India. Population dynamics of three cosmopolitan inversions has been studied in Indian population of D. ananassae and it is evident from the results that there is a considerable degree of genetic divergence at the level of inversion polymorphism. In general, the populations from south India show more differentiation than those from the north. These three cosmopolitan inversions, which are coextensive with the species, exhibit heterosis. Interracial hybridization does not lead to beaakdown of heterosis, which suggests that evidence for coadaptation is lacking in geographic populations of D. ananassae. Heterosis appears to be simple luxuriance rather than populational heterosis (coadaptation). Unlinked inversions occur in random associations, indicating no interchromosomal interactions. However, two inversions of the third chromosome often show strong linkage disequilibrium in laboratory populations, which is due to epistatic gene interaction and suppression of crossing-over. Genetic variations for certain allozyme polymorphism and sternoleural bristle phenotypes in Indian populations of D. ananassae have also been observed.A number of investigations have also been carried out on certain aspects of behaviour genetics of Indian D. ananassae. There is evidence for sexual isolation within D. ananassae. Significant variations in mating propensity of several isofemale strains, inversion karyotypes, the diminishing effects of certain mutations on sexual activity of males and positive response to selection for high and low mating propensity provide evidence for genetic control of sexual behaviour in D. ananassae. Males contribute more to variation and thus are more subject to intra-sexual selection than females. Evidence for rare male mating advantage has also been presented. Geographic strains of D. ananassae show variation with respect to oviposition site preference. The results of studies on pupation site preference, which is an important component of larval behaviour, suggest that larval pupation behaviour in D. ananassae is under polygenic control with a substantial amount of additive genetic variation.     

A framework for the study of genetic variation in migratory behaviour

Evolutionary change results from selection acting on genetic variation. For migration to be successful, many different aspects of an animal’s physiology and behaviour need to function in a co-coordinated way. Changes in one migratory trait are therefore likely to be accompanied by changes in other migratory and life-history traits. At present, we have some knowledge of the pressures that operate at the various stages of migration, but we know very little about the extent of genetic variation in various aspects of the migratory syndrome. As a consequence, our ability to predict which species is capable of what kind of evolutionary change, and at which rate, is limited. Here, we review how our evolutionary understanding of migration may benefit from taking a quantitative-genetic approach and present a framework for studying the causes of phenotypic variation. We review past research, that has mainly studied single migratory traits in captive birds, and discuss how this work could be extended to study genetic variation in the wild and to account for genetic correlations and correlated selection. In the future, reaction-norm approaches may become very important, as they allow the study of genetic and environmental effects on phenotypic expression within a single framework, as well as of their interactions. We advocate making more use of repeated measurements on single individuals to study the causes of among-individual variation in the wild, as they are easier to obtain than data on relatives and can provide valuable information for identifying and selecting traits. This approach will be particularly informative if it involves systematic testing of individuals under different environmental conditions. We propose extending this research agenda by using optimality models to predict levels of variation and covariation among traits and constraints. This may help us to select traits in which we might expect genetic variation, and to identify the most informative environmental axes. We also recommend an expansion of the passerine model, as this model does not apply to birds, like geese, where cultural transmission of spatio-temporal information is an important determinant of migration patterns and their variation.     

Welfare of sheep: Providing for welfare in an extensive environment

As a species traditionally managed extensively, at least for some of the year, sheep have received relatively little attention from a welfare perspective. Although extensively managed animals have greater behavioural freedom than those managed intensively, they are still vulnerable to other welfare challenges. Welfare can be considered from the perspective of the animals biological functioning, the naturalness of the way in which they are kept, or their feelings (negative or positive). These different domains can be integrated by considering the animal (and the adaptations it possesses) and the environment (made up of the challenges the animal experiences). In this schema, the extensively managed animal may experience poor welfare if the environmental challenges overwhelm its evolved coping strategies (for example, in high temperatures if behavioural and physiological adaptations cannot dissipate heat sufficiently) or if it has adaptations (such as behavioural anti-predator responses) that are no longer required. To assess welfare of sheep, therefore, we need to know when the animal may begin to suffer if its adaptations do not meet the challenges presented by the environment or when the cost of meeting those challenges is too great. In addition, the adaptations the sheep possess will affect how well it copes, e.g. breed differences in maternal behaviour and responses to predators will influence lamb survival and predation risk in extensive environments, respectively. How well the extensive environment can meet the behavioural needs of sheep has never been tested. However, consideration of the behaviour and habitat of wild sheep suggests that some preferred environmental features, such as escape terrain or specific birth sites, may not be present in farmed environments. How this affects sheep welfare is still to be determined. Sheep welfare is also affected by the quality of the stockperson caring for them. Interactions with people for extensively managed sheep are infrequent, but nearly always aversive, and whether the nature of interactions can be affected by stockperson attitude and behaviour, as seen in intensively managed species, remains to be seen. There is evidence, however, that farmers may under-estimate the negative impact that themselves and their sheepdogs can have on their sheep. Farmer attitude can also affect management decisions which may have an indirect impact on the welfare of their sheep. For example, farmer attitudes towards lameness are related to treatment decisions. The extensive environment thus provides particular challenges to welfare that differ markedly from those in intensive systems, and still need further investigation.     

羚牛生态生物学研究现状

羚牛是介于羊和牛之间分类地位特殊的偶蹄目牛科动物, 共有4 个亚种。中国是羚牛的主要分布区, 拥有世界上绝大多数的羚牛种群。有关羚牛的研究主要由中国的科研人员在中国境内完成。羚牛的生态生物学研究主要涉及到地理分布、种群数量、栖息地、季节性迁移活动、集群行为、家域、日活动规律、采食习性、繁殖和防御行为。对羚牛研究现状的综述性介绍即从这10 个方面进行。     

Life of Listeria monocytogenes in the host cells' cytosol

In the past decades impressive knowledge has been accumulated concerning the basic mechanisms of interactions between intracellular bacteria and their host cells. Comparatively little is known on the metabolic requirements necessary for efficient replication of these bacteria within their specific host cell compartments. Recent developments in functional genomics have led to more extensive studies of the metabolic aspects that may be crucial for understanding the pathogenesis of intracellular bacteria. Here we summarize our present knowledge on the physiology of L. monocytogenes with emphasis on those parts that seem to be important for its ability to replicate in the cytosol of mammalian host cells.     

Sheep model in orthopedic research: a literature review

The aim of the study reported here was to provide some basic and general information on the suitability of an experimental sheep model for conducting in vivo orthopedic studies. The authors have classified the fundamental aspects that should be carefully evaluated when using sheep as an experimental model in orthopedic research: factors strictly related to bone anatomy and formation; and factors strictly affecting bone physiology, such as gastrointestinal mineral and vitamin absorption, and reproductive cycle. Future investigations should address all of the aspects highlighted, since there is no animal with the same anatomic, biochemical, physiologic, and biological characteristics as those of human beings. Moreover, useful data for treating orthopedic patients are based not only on good planning and study design, but also on perfect knowledge of the animal used and the differences between the model and the human being. The authors hope that this report will contribute to extrapolation of reliable data for use of sheep in the orthopedics field.     

The evolutionary ecology of alternative migratory tactics in salmonid fishes

Extensive individual variation in spatial behaviour is a common feature among species that exhibit migratory life cycles. Nowhere is this more evident than in salmonid fishes; individual fish may complete their entire life cycle in freshwater streams, others may migrate variable distances at sea and yet others limit their migrations to larger rivers or lakes before returning to freshwater streams to spawn. This review presents evidence that individual variation in migratory behaviour and physiology in salmonid fishes is controlled by developmental thresholds and that part of the variation in proximal traits activating the development of alternative migratory tactics is genetically based. We summarize evidence that alternative migratory tactics co‐exist within populations and that all individuals may potentially adopt any of the alternative phenotypes. Even though intra‐specific genetic divergence of migratory tactics is uncommon, it may occur if female competition for oviposition sites results in spawning segregation of alternative phenotypes. Because of their polygenic nature, alternative migratory tactics are considered as threshold traits. Threshold traits have two characteristics: an underlying 'liability' trait that varies in a continuous fashion, and a threshold value which is responsible for the discreetness observed in phenotypic distribution. We review evidence demonstrating that body size is an adequate proxy for the liability trait controlling the decision to migrate, but that the same phenotypic outcome (anadromy or residency) may be reached by different developmental pathways. The evidence suggesting a significant heritable component in the development of alternative migratory tactics is subsequently reviewed, leading us to conclude that alternative migratory tactics have considerable potential to respond to selection and evolve. We review what is known about the proximal physiological mechanisms mediating the translation of the continuous value of the liability trait into a discontinuous migratory tactic. We conclude by identifying several avenues for future research, including testing the frequency‐dependent selection hypothesis, establishing the relative importance of adaptive phenotypic plasticity in explaining some geographic gradients in migratory behaviour and identifying the physiological and genetic basis of the switching mechanisms responsible for alternative migratory tactics.     

The function of granulocytes stored in whole blood in relation to the storage time

Whereas numerous investigations and extensive experiences have been available for some years on problems of storing erythrocytes and thrombocytes, it was only recently that the storage of granulocytes with the aim at maintaining their function for a longer period has been given proper regard as a problem, which, however, could not be brought to a satisfactory solution. Primarily, attention is focused on specific cell physiology. At the same time it is necessary to have at hand a number of approaches covering as many partial aspects of the granulocyte function as possible to identify the functional damages or the extent of cell damage in connection with storage. In the present paper the behaviour of granulocytes under conditions commonly used in transfusion medicine for storing blood are investigated for a period of 4 days by controlling the total leukocyte and granulocyte values, pH value, osmolality, vitality test in the colour excluding test as well as by function tests such as behaviour of phagocytosis and NBT capability of reduction. A report is presented on the results of quantitative and qualitative changes in the investigated parameters.     

Alternative life-history styles of some African cichlid fishes

Synopsis About 50% of African cichlid species have been described. Little is known about cichlid ecology, behaviour or about the evolution and interactions of communities. Nevertheless, trends which provide an insight into cichlid life histories, into evolutionary alternatives and into behavioural alternatives which may be followed during the life time of these fishes are emerging. Cichlids which spend their entire life history in a single habitat belong to species flocks that have spectated greatly. In contrast, those groups which live in a variety of habitats during their life history have spectated little. Despite the trophic specializations that have occurred among cichlids, many and perhaps all species, have the ability to feed upon alternative food resources. They do so by switching or by markedly modifying their behaviour and all are particularly opportunistic. The extensive adaptive radiation of cichlids with regard to those attributes of morphology and behaviour that are associated with survival and growth, is apparently not matched by similar radiations in reproductive behaviour. The perceived evolutionary conservatism with regard to reproductive behaviour is attributed to the effects of stabilizing selection, but this still needs to be tested. The constraints of stabilizing selection appear to be lifted with regard to coloration. If colour is an important component of the specific-mate recognition system then it should also be subjected to stabilizing selection so the readiness with which colour variation occurs within and between populations needs explanation. Parental care in cichlids has followed a variety of evolutionary alternatives, all of which are geared to improve the chances of survival of offspring in their specific micro-habitats.     

Molecular population genetics and the search for adaptive evolution in plants

The first papers on plant molecular population genetics were published approximately 10 years ago. Since that time, well over 50 additional studies of plant nucleotide polymorphism have been published, and many of these studies focused on detecting the signature of balancing or positive selection at a locus. In this review, we discuss some of the theoretical and statistical issues surrounding the detection of selection, with focus on plant populations, and we also summarize the empirical plant molecular population genetics literature. At face value, the literature suggests that a history of balancing or positive selection in plant genes is rampant. In two well-studied taxa (maize and Arabidopsis) over 20% of studied genes have been interpreted as containing the signature of selection. We argue that this is probably an overstatement of the prevalence of natural selection in plant genomes, for two reasons. First, demographic effects are difficult to incorporate and have generally not been well integrated into the plant population genetics literature. Second, the genes studied to date are not a random sample, so selected genes may be overrepresented. The next generation of studies in plant molecular population genetics requires additional sampling of local populations, explicit comparisons among loci, and improved theoretical methods to control for demography. Eventually, candidate loci should be confirmed by explicit consideration of phenotypic effects.     

Physiological effects of dominance hierarchies: laboratory artefacts or natural phenomena?

Studies of fish behaviour have demonstrated the existence of social interactions that result in dominance hierarchies. In environments in which resources, such as food, shelter and mates, are limited, social competition results in some fish becoming dominant and occupying the most profitable positions. This behaviour has been observed in natural environments and also in many laboratory‐based experiments. When two fish have been confined in a small tank, one of them usually has exhibited behaviour that suggests it is dominant over the other submissive animal. Physiological consequences of social interaction can be seen in both dominants and subordinates but are more extreme in the subordinate. However, this scenario is without doubt an artificial situation. Fewer experiments have been conducted using laboratory experiments that are more socially and physically complex than those experienced by dyads in tanks. In simple fluvial tanks, through which water is recirculated, the physiological responses of fish to social competition have generally been qualitatively similar to those recorded among dyads. However, when environmental disturbances, complex resource distributions, increase in water flushing, presence of predators and competing species of fish have been included in experimen‐tal designs, there have been fewer, diminished or no physiological dierences between dominant and subordinate fish. There have been very few studies of physiology in relation to dominance in natural habitats, and those that have been conducted suggest that under some circumstances hierarchies may cause less intense physiological responses than have been suggested based on results of laboratory studies in simple environments. Possible reasons for these variations are discussed. The need is identified for a well structured experimental approach to the investi‐gation of the causes and consequences of hierarchies if the ecology of wild fish is to be modelled eectively based on physiological processes. It is also suggested that the further development and application of techniques for monitoring physiologies of fish in the wild is important.     

GABAA receptor subtype involvement in addictive behaviour

GABA_A receptors form the major class of inhibitory neurotransmitter receptors in the mammalian brain. This review sets out to summarize the evidence that variations in genes encoding GABA_A receptor isoforms are associated with aspects of addictive behaviour in humans, while animal models of addictive behaviour also implicate certain subtypes of GABA_A receptor. In addition to outlining the evidence for the involvement of specific subtypes in addiction, we summarize the particular contributions of these isoforms in control over the functioning of brain circuits, especially the mesolimbic system, and make a first attempt to bring together evidence from several fields to understanding potential involvement of GABA_A receptor subtypes in addictive behaviour. While the weight of the published literature is on alcohol dependency, the underlying principles outlined are relevant across a number of different aspects of addictive behaviour.     

Rethinking phenotypic plasticity and its consequences for individuals,populations and species

Much research has been devoted to identify the conditions under which selection favours flexible individuals or genotypes that are able to modify their growth, development and behaviour in response to environmental cues, to unravel the mechanisms of plasticity and to explore its influence on patterns of diversity among individuals, populations and species. The consequences of developmental plasticity and phenotypic flexibility for the performance and ecological success of populations and species have attracted a comparatively limited but currently growing interest. Here, I re-emphasize that an increased understanding of the roles of plasticity in these contexts requires a ‘whole organism'' (rather than ‘single trait'') approach, taking into consideration that organisms are integrated complex phenotypes. I further argue that plasticity and genetic polymorphism should be analysed and discussed within a common framework. I summarize predictions from theory on how phenotypic variation stemming from developmental plasticity and phenotypic flexibility may affect different aspects of population-level performance. I argue that it is important to distinguish between effects associated with greater interindividual phenotypic variation resulting from plasticity, and effects mediated by variation among individuals in the capacity to express plasticity and flexibility as such. Finally, I claim that rigorous testing of predictions requires methods that allow for quantifying and comparing whole organism plasticity, as well as the ability to experimentally manipulate the level of and capacity for developmental plasticity and phenotypic flexibility independent of genetic variation.     

Lizards as model organisms for linking phylogeographic and speciation studies

Lizards have been model organisms for ecological and evolutionary studies from individual to community levels at multiple spatial and temporal scales. Here we highlight lizards as models for phylogeographic studies, review the published population genetics/phylogeography literature to summarize general patterns and trends and describe some studies that have contributed to conceptual advances. Our review includes 426 references and 452 case studies: this literature reflects a general trend of exponential growth associated with the theoretical and empirical expansions of the discipline. We describe recent lizard studies that have contributed to advances in understanding of several aspects of phylogeography, emphasize some linkages between phylogeography and speciation and suggest ways to expand phylogeographic studies to test alternative pattern‐based modes of speciation. Allopatric speciation patterns can be tested by phylogeographic approaches if these are designed to discriminate among four alternatives based on the role of selection in driving divergence between populations, including: (i) passive divergence by genetic drift; (ii) adaptive divergence by natural selection (niche conservatism or ecological speciation); and (iii) socially‐mediated speciation. Here we propose an expanded approach to compare patterns of variation in phylogeographic data sets that, when coupled with morphological and environmental data, can be used to to discriminate among these alternative speciation patterns. [Correction made after online publication (28/07/2010): (minor deletion in the last line of the abstract)].     

Genetic improvement of mammalian fertility: A review of opportunities

The possibility of genetic improvement in the reproductive performance of domestic mammals is introduced. The paper is principally concerned with biological variation and change, but the need to consider economic factors when relating change to improvement is illustrated from time to time. Both male traits, such as semen quality, and female traits, such as litter size, are considered.There is considerable variation among existing populations and the importance of the choice of the most suitable of available populations whether pure or crossbred is emphasized. Variation in litter size is greatest among breeds of sheep (approx. 300%) and least among cattle (only 4%). The possibility that differences in performance may be specific to particular environments and the implications of such an interaction between genotype and environment for the choice of populations are discussed. Among cattle, variation in the incidence of calving difficulties may be marked; it may increase three-fold in cows mated to bulls of a large breed relative to cows mated to bulls of a small breed. Cross-breeding may improve reproductivity by 20%.Once the best population has been chosen, further improvement is dependent on selection within that population. Selection within populations has been rare due to low predicted rates of response. Low selection differentials have probably contributed more to slow responses than have low heritabilities, and recent marked changes achieved by group breeding schemes illustrate what can be done with intense selection.A principal reason for small selection differentials is the inability to measure all reproductive traits in both sexes; one sex has to be chosen at random. The common physiology of reproduction in males and females indicates common genetic control, and the possibility of indirect selection on, say, testis growth to alter ovarian activity is considered along with the use of other physiological selection criteria, including ovulation rate and hormone levels, to increase the rate of response to selection.Finally, the implementation of improvement schemes is discussed. The improvement of reproduction has to be balanced against improvement in other traits such as growth; as the benefits of selection for other traits decline, reproduction would receive greater emphasis. International collaboration to develop strains of extremely high merit for subsequent use by crossbreeding is advocated.     

Animal Board Invited Review: Sheep birth distribution in past herds: a review for prehistoric Europe (6th to 3rd millennia BC)

In temperate latitudes sheep have a seasonal reproductive behaviour, which imposes strong constraints on husbandry in terms of work organization and availability of animal products. During the last 50 years, researchers have focused on understanding the mechanisms driving small ruminants’ reproduction cycles and finding ways to control them. This characteristic is inherited from their wild ancestor. However, the history of its evolution over the 10 millennia that separates present day European sheep from their Near Eastern ancestors’ remains to be written. This perspective echoes archaeologists’ current attempts at reconstructing ancient pastoral societies’ socio-economical organization. Information related to birth seasonality may be retrieved directly from archaeological sheep teeth. The methodology consists of reconstructing the seasonal cycle record in sheep molars, through sequential analysis of the stable oxygen isotope composition (δ¹⁸O) of enamel. Because the timing of tooth development is fixed within a species, inter-individual variability in this parameter reflects birth seasonality. A review of the data obtained from 10 European archaeological sites dated from the 6th to the 3rd millennia BC is provided. The results demonstrate a restricted breeding season for sheep: births occurred over a period of 3 to 4 months, from late winter to early summer at latitudes 43°N to 48°N, while a later onset was observed at a higher latitude (59°N). All conclusions concurred with currently held expectations based on present day sheep physiology, which, aside from the historical significance, contributes to the reinforcing of the methodological basis of the approach. Further study in this area will permit regional variability attributable to technical choices, within global schemes, to be fully reported.