Combining hydrodynamic modelling with genetics: can passive larval drift shape the genetic structure of Baltic Mytilus populations?

While secondary contact between Mytilus edulis and Mytilus trossulus in North America results in mosaic hybrid zone formation, both species form a hybrid swarm in the Baltic. Despite pervasive gene flow, Baltic Mytilus species maintain substantial genetic and phenotypic differentiation. Exploring mechanisms underlying the contrasting genetic composition in Baltic Mytilus species will allow insights into processes such as speciation or adaptation to extremely low salinity. Previous studies in the Baltic indicated that only weak interspecific reproductive barriers exist and discussed the putative role of adaptation to environmental conditions. Using a combination of hydrodynamic modelling and multilocus genotyping, we investigate how oceanographic conditions influence passive larval dispersal and hybrid swarm formation in the Baltic. By combining our analyses with previous knowledge, we show a genetic transition of Baltic Mytilus species along longitude 12°‐13°E, that is a virtual line between Malmö (Sweden) and Stralsund (Germany). Although larval transport only occurs over short distances (10–30 km), limited larval dispersal could not explain the position of this genetic transition zone. Instead, the genetic transition zone is located at the area of maximum salinity change (15–10 psu). Thus, we argue that selection results in weak reproductive barriers and local adaptation. This scenario could maintain genetic and phenotypic differences between Baltic Mytilus species despite pervasive introgressive hybridization.     

The extracellular matrix: an active or passive player in fibrosis?

Fibrosis is characterized by excessive accumulation of collagen and other extracellular matrix (ECM) components, and this process has been likened to aberrant wound healing. The early phases of wound healing involve the formation of a provisional ECM containing fibrin, fibrinogen, and fibronectin. Fibroblasts occupy this matrix and proliferate in response to activators elaborated by leukocytes that have migrated into the wound and are retained by the ECM. This coincides with the appearance of the myofibroblast, a specialized form of fibroblast whose differentiation is primarily driven by cytokines, such as transforming growth factor-β (TGF-β), and by mechanical tension. When these signals are reduced, as when TGF-β secretion is reduced, or as in scar shrinkage, myofibroblasts undergo apoptosis, resulting in a collagen-rich, cell-poor scar. Retention of myofibroblasts in fibrosis has been described as the result of imbalanced cytokine signaling, especially with respect to levels of activated TGF-β. ECM components can regulate myofibroblast persistence directly, since this phenotype is dependent on extracellular hyaluronan, tenascin-C, and the fibronectin splice variant containing the "extra domain A," and also, indirectly, through retention of TGF-β-secreting cells such as eosinophils. Thus the ECM is actively involved in both cellular and extracellular events that lead to fibrosis. Targeting components of the ECM as cells respond to injury and inflammatory stimuli holds promise as a means to avoid development of fibrosis and direct the wound-healing process toward reestablishment of a healthy equilibrium.     

Maternal inheritance and rapid evolution of sexual size dimorphism: passive effects or active strategies?

Adaptive evolution is often strongly influenced by maternal inheritance that transfers the parental strategies across generations. The consequences of maternal effects for the offspring generation depend on the between-generation similarity in environments and on the evolved sensitivity of the offspring's ontogeny to maternal effects. When these factors differ between sons and daughters, maternal effects can influence the evolution of sexual dimorphism. The establishment of house finch populations across western Montana during the last 30 years was accompanied by rapid evolutionary change in sexual size dimorphism. Here I show that traits that changed the most across generations were most influenced by maternal effects in males but not females. Maternal effects differentially affected sons' and daughters' survival; greater maternal effects were commonly associated with higher survival of sons, especially when maternal and offspring environments were similar. Stronger maternal effects extended preselection phenotypic variance in morphological traits of males, thereby producing some locally adaptive phenotypes and lessening juvenile mortality. Thus, the observed sex-specific maternal effects and their contribution to the evolution of sexual size dimorphism are likely a passive consequence of the distinct sensitivity of sons and daughters to maternal adaptations to breeding in ecologically distinct parts of the house finch's expanding range.     

Character displacement: in situ evolution of novel phenotypes or sorting of pre‐existing variation?

Character displacement - the divergence of traits between species in response to competition for resources or mates - has long been viewed as a major cause of adaptive diversification and species coexistence. Yet, we lack answers to basic questions concerning the causes and consequences of character displacement, not the least of which is why some species are more prone than others to undergo character displacement. Here, we address these questions by describing how character displacement can proceed through two nonexclusive routes that differ in the source of phenotypic variation, and, hence, in the ease with which character displacement may unfold. During in situ evolution of novel phenotypes, new traits that are divergent from a heterospecific competitor are generated and spread in sympatry. During sorting of pre-existing variation, such traits are initially favoured in allopatry before the two species encounter one another. Later, when they come into contact, character displacement transpires when these pre-existing divergent phenotypes increase in frequency in sympatry relative to allopatry. Because such sorting of pre-existing variation should unfold relatively rapidly, we suggest that species that express resource or mating polymorphism prior to interactions with heterospecifics may be more prone to undergo character displacement. We discuss the key differences between these two routes, review possible examples of each, and describe how the distinction between them provides unique insights into the evolutionary consequences of species interactions, the origins of diversity, and the factors that govern species coexistence.     

The antigenic evolution of influenza: drift or thrift?

It is commonly assumed that antibody responses against the influenza virus are polarized in the following manner: strong antibody responses are directed at highly variable antigenic epitopes, which consequently undergo ‘antigenic drift’, while weak antibody responses develop against conserved epitopes. As the highly variable epitopes are in a constant state of flux, current antibody-based vaccine strategies are focused on the conserved epitopes in the expectation that they will provide some level of clinical protection after appropriate boosting. Here, we use a theoretical model to suggest the existence of epitopes of low variability, which elicit a high degree of both clinical and transmission-blocking immunity. We show that several epidemiological features of influenza and its serological and molecular profiles are consistent with this model of ‘antigenic thrift’, and that identifying the protective epitopes of low variability predicted by this model could offer a more viable alternative to regularly update the influenza vaccine than exploiting responses to weakly immunogenic conserved regions.     

Repeated antipredator conditioning: a pathway to habituation or to better avoidance?

Hatchery‐reared juveniles of Arctic charr Salvelinus alpinus were conditioned to the odours of Arctic charr‐fed pikeperch Sander lucioperca in the absence of any other cues. Accordingly, there was no physical threat of capture for the Arctic charr. It was evident from the subsequent survival tests that a single exposure to predator odours was enough to increase Arctic charr survival compared to predator‐naïve control fish whist under direct threat from live predators. Instead of habituating to predator odours, the fish conditioned repeatedly (four times) improved their spatial avoidance of predator cues in the course of training. The repeated conditioning also further enhanced the survival of the test fish as compared to the singly conditioned fish. The economical and ethical advantages of training with chemical cues, combined with its high reliability, could promote the success of fish reintroductions especially through repeated antipredator conditioning.     

The role of sleep in declarative memory consolidation: passive,permissive, active or none?

Those inclined to relish in scientific controversy will not be disappointed by the literature on the effects of sleep on memory. Opinions abound. Yet refinements in the experimental study of these complex processes of sleep and memory are bringing this fascinating relationship into sharper focus. A longstanding position contends that sleep passively protects memories by temporarily sheltering them from interference, thus providing precious little benefit for memory. But recent evidence is unmasking a more substantial and long-lasting benefit of sleep for declarative memories. Although the precise causal mechanisms within sleep that result in memory consolidation remain elusive, recent evidence leads us to conclude that unique neurobiological processes within sleep actively enhance declarative memories.     

Between‐year variation of MHC allele frequencies in great reed warblers: selection or drift?

The major histocompatibility complex (MHC) genes are extremely polymorphic and this variation is assumed to be maintained by balancing selection. Cyclic interactions between pathogens and their hosts could generate such selection, and specific MHC alleles or heterozygosity at certain MHC loci have been shown to confer resistance against particular pathogens. Here we compare the temporal variation in allele frequencies of 23 MHC class I alleles with that of 23 neutral microsatellite markers in adult great reed warblers (a passerine bird) in nine successive cohorts. Overall, the MHC alleles showed a significantly higher variation in allele frequencies between cohorts than the microsatellite alleles, using a multi-variate genetic analysis (amova). The frequency of two specific MHC alleles, A3e (P = 0.046) and B4b (P = 0.0018), varied more between cohorts than expected from random, whereas none of the microsatellite alleles showed fluctuations exceeding the expectation from stochastic variation. These results imply that the variation in MHC allele frequencies between cohorts is not a result of demographic events, but rather an effect of selection favouring different MHC alleles in different years.     

Always together: mate guarding or predator avoidance as determinants of group cohesion in white‐breasted mesites?

Being a member of a cohesive group can have fitness benefits such as decreased predation risk, increased feeding efficiency as well as enhanced access to social information and mates. However, competition and the risk of parasite transmission exert centrifugal forces on group‐living. Thus, the actual degree of cohesion is expected to vary as a function of the relative importance of several social and ecological factors. White‐breasted mesites Mesitornis variegata are medium‐sized ground‐dwelling birds endemic to the dry deciduous forests of western Madagascar. They live in stable breeding pairs or small family groups, mate monogamously and often form temporary heterospecific associations with canopy‐dwelling bird species that give alarm calls to which mesites respond with anti‐predator behaviours. We investigated the potential effects of predation risk and mate defence on mesite group cohesion by analysing inter‐individual distances of 20 groups as a function of mesite social organization, alarm call events, the size of associated heterospecific flocks, and the adults' reproductive state. Mesite social units were very cohesive, particularly in families, when associated with smaller heterospecific flocks, and after an alarm call event. Adult reproductive state did not influence breeding partners' cohesion. Therefore, the pronounced group cohesion in mesites seems to be mainly a response to the high predation risk typically associated with a terrestrial life‐style, and not to mate‐guarding. However, we suggest that high group cohesion due to predation risk could limit opportunities for solitary extra‐territorial forays to obtain extra‐pair copulations, thereby contributing to a strictly monogamous system in this species.     

Behavioral responses of native prey to disparate predators: naiveté and predator recognition

It is widely accepted that predator recognition and avoidance are important behaviors in allowing prey to mitigate the impacts of their predators. However, while prey species generally develop anti-predator behaviors through coevolution with predators, they sometimes show accelerated adoption of these behaviors under strong selection pressure from novel species. We used a field manipulation experiment to gauge the ability of the common ringtail possum (Pseudocheirus peregrinus), a semi-arboreal Australian marsupial, to recognize and respond to olfactory cues of different predator archetypes. We predicted that ringtails would display stronger anti-predator behaviors to cues of the invasive European red fox (Vulpes vulpes) in areas where fox impacts had been greatest, and to cues of the native lace monitor (Varanus varius) in areas of sympatry compared with allopatry. We found that ringtails fled quickly and were more alert when exposed to the fecal odors of both predators compared to neutral and pungent control odors, confirming that predator odors are recognized and avoided. However, these aversive responses were similar irrespective of predator presence or level of impact. These results suggest that selection pressure from the fox has been sufficient for ringtails to develop anti-predator behaviors over the few generations since foxes have become established. In contrast, we speculate that aversive responses by ringtails to the lace monitor in areas where this predator is absent reflect recent coexistence of the two species. We conclude that rapid evolution of anti-predator behaviors may occur when selection is strong. The maintenance of these behaviors should allow re-establishment of predator–prey relationships if the interactants regain sympatry via range shifts or management actions to reintroduce them to their former ranges.     

Human T-cell responses to malaria: mostly forgotten or committed to memory?

T cells have been implicated in both malaria immunity and malaria disease and factors controlling the maintenance of T-cell responses over time may alter the clinical outcome o f infection. Michael Good and Janine Bilsborough have compared the T-cell responses to epitopes from the Plasmodium vivax and P. falciparum circumsporozoite proteins and here discuss the issue of T-cell memory as it applies to malaria.     

Localized cellular responses to Ichthyophthirius multifiliis: protection or pathogenesis?

The ciliate Ichthyophthirius multifiliis is an economically important parasite in aquaculture as well as in the ornamental fish trade. The parasite has been well studied and details of life cycle, host response and pathogenesis have been characterized. Parasite development within host epithelial tissues initiates localized leukocytic infiltrations, although the relationship between these responses and host resistance is uncertain, and whether or not leukocyte responses play a role in protective immunity is unclear. Here Frank Cross outlines the character of localized cellular responses during I. multifiliis infection and discusses the contribution of such responses to protective immunity and pathogenesis.     

Trade-offs between the reproductive and immune systems: facultative responses to resources or obligate responses to reproduction?

A major challenge in biology is understanding how organisms partition limited resources among physiological processes. For example, offspring production and self-maintenance are important for fitness and survival, yet these critical processes often compete for resources. While physiological trade-offs between reproduction and immune function have been documented, their regulation remains unclear. Most current evidence suggests that physiological changes during specific reproductive states directly suppress various components of the immune system; however, some studies have not found this clear relationship. We performed two experiments in female tree lizards (Urosaurus ornatus) that demonstrate the presence of trade-offs between the reproductive and immune systems under controlled laboratory conditions. These results also support the hypothesis that these trade-offs are a facultative response to resource availability and are not obligatory responses to physiological changes during reproduction. We found that (1) experimentally increasing reproductive investment under limited resources resulted in suppressed immune function and (2) experimentally limiting resources resulted in immunosuppression but only during resource costly reproductive activities. There seems to be a critical balance of resources that is maintained between multiple processes, and changes in the balance between energy intake and output can have major consequences for immune function.     

Endocrine responses to unpredictable environmental events: stress or anti-stress hormones?

In addition to seasonal changes in morphology, physiology andbehavior that occur in predictable annual cycles, there arefacultative responses to unpredictable events known as labile(i.e., short-lived) perturbation factors (LPFs). These rapidbehavioral and physiological changes have been termed the "emergency"life history stage (ELHS) and serve to enhance life-time fitness.Glucocorticosteroids interacting with other hormones in thehypothalamo-pituitary-adrenal (HPA) cascade, initiate and orchestratethe ELHS within minutes to hours. Components of the ELHS include:redirection of behavior from a normal life history stage toincreased foraging, irruptive-type migration during the day,enhanced restfulness at night, elevated gluconeogenesis andrecovery once the perturbation passes. These physiological andbehavioral changes allow an individual to avoid potential deleteriouseffects of stress that may result from chronically elevatedlevels of circulating glucocorticosteroids over days and weeks.In other words, acute rises in glucocorticosteroids followingperturbations of the environment may actually avoid chronicstress and serve primarily as "anti-stress" hormones. Severalfield studies in diverse habitats indicate that free-livingpopulations have elevated circulating levels of corticosteroidswhen in an ELHS. However, expression of an ELHS may not alwaysbe advantageous and there is accumulating evidence from birdsthat the adrenocortical responses to LPFs are modulated bothon seasonal and individual levels. These data suggest that glucocorticosteroidsecretions in response to LPFs not only trigger physiologicaland behavioral responses but also allow flexibility so thatthe response is integrated in relation to time of year (normalLHS) as well as individual differences owing to body condition,disease and social status.     

Do population indicators work? Investigating correlated responses of bird populations in relation to predator management

Using population indicators to evaluate conservation achievements is widely practised, yet seldom empirically tested. If populations are consistently correlated in response to a shared ecological driver, the indicator species approach can be used as a cost-effective, ecologically-based shortcut to measuring the effects of conservation management. Long-term monitoring of forest bird populations associated with mammalian pest control programmes in New Zealand provides a useful framework for testing the population indicator species concept. We evaluated population trends in 21 bird species vulnerable to predation by introduced mammals (primarily mustelids and rodents) at managed and unmanaged beech (Nothofagus) forest sites. Correlated population trends between species pairs were detected at individual sites. However, neither positive nor negative correlations in species trends could be predicted by life history traits and predator management did not produce consistent, correlated population trends among sites. Our results do not support the use of a population indicator approach to management and reporting for forest birds in New Zealand. Relationships between purported indicator taxa and other species need to be understood for various management scenarios before population indicators can be confidently applied to measuring conservation achievement.