Novel Venom Proteins Produced by Differential Domain-Expression Strategies in Beaded Lizards and Gila Monsters (genus Heloderma)

The origin and evolution of venom proteins in helodermatid lizards were investigated by multidisciplinary techniques. Our analyses elucidated novel toxin types resultant from three unique domain-expression processes: 1) The first full-length sequences of lethal toxin isoforms (helofensins) revealed this toxin type to be constructed by an ancestral monodomain, monoproduct gene (beta-defensin) that underwent three tandem domain duplications to encode a tetradomain, monoproduct with a possible novel protein fold; 2) an ancestral monodomain gene (encoding a natriuretic peptide) was medially extended to become a pentadomain, pentaproduct through the additional encoding of four tandemly repeated proline-rich peptides (helokinestatins), with the five discrete peptides liberated from each other by posttranslational proteolysis; and 3) an ancestral multidomain, multiproduct gene belonging to the vasoactive intestinal peptide (VIP)/glucagon family being mutated to encode for a monodomain, monoproduct (exendins) followed by duplication and diversification into two variant classes (exendins 1 and 2 and exendins 3 and 4). Bioactivity characterization of exendin and helokinestatin elucidated variable cardioactivity between isoforms within each class. These results highlight the importance of utilizing evolutionary-based search strategies for biodiscovery and the virtually unexplored potential of lizard venoms in drug design and discovery.

Lacustrine evidence of Holocene environmental change from three Faroese lakes: a multiproxy XRF and stable isotope study

The vegetation history of the Faroe Islands has been investigated in numerous studies all broadly showing that the early-Holocene vegetation of the islands largely consisted of fellfield with gravely and rocky soils formed under a continental climate which shifted to an oceanic climate around 10,000 cal yr BP when grasses, sedges and finally shrubs began to dominant the islands. Here we present data from three lake sediment cores and show a much more detailed history from geochemical and isotope data. These data show that the Faroe Islands were deglaciated by the end of Younger Dryas (11,700 10,300 cal yr BP), at this time relatively high sedimentation rates with high delta C-13 imply poor soil development. delta C-13, Ti and chi data reveal a much more stable and warm mid-Holocene until 7410 cal yr BP characterised by increasing vegetation cover and build up of organic soils towards the Holocene thermal maximum around 7400 cal yr BP. The final meltdown of the Laurentide ice sheet around 7000 cal yr BP appears to have impacted both ocean and atmospheric circulation towards colder conditions on the Faroe Islands. This is inferred by enhanced weathering and increased deposition of surplus sulphur (sea spray) and erosion in the highland lakes from about 7400 cal yr BP. From 4190 cal yr BP further cooling is believed to have occurred as a consequence for increased soil erosion due to freeze/thaw sequences related to oceanic and atmospheric variability. This cooling trend appears to have advanced further from 3000 cal yr BR A short period around 1800 cal yr BP appears as a short warm and wet phase in between a general cooling characterised by significant soil erosion lasting until 725 cal yr BP. Interestingly, increased soil erosion seems to have begun at 1360 cal yr BP, thus significantly before the arrival of the first settlers on the Faroe Island around 1150 cal yr BP, although additional erosion took place around 1200 cal yr BP possibly as a consequence of human activities. Hence it appears that if humans caused a change in the Faroe landscape in terms of erosion they in fact accelerated a process that had already started. Soil erosion was a dominant landscape factor during the Little Ice Age, but climate related triggers can hardly be distinguished from human activities. (c) 2010 Elsevier Ltd. All rights reserved.

Fitness costs associated with mounting a social immune response

Social immune systems comprise immune defences mounted by individuals for the benefit of others (sensu Cotter & Kilner 2010a). Just as with other forms of immunity, mounting a social immune response is expected to be costly but so far these fitness costs are unknown. We measured the costs of social immunity in a sub-social burying beetle, a species in which two or more adults defend a carrion breeding resource for their young by smearing the flesh with antibacterial anal exudates. Our experiments on widowed females reveal that a bacterial challenge to the breeding resource upregulates the antibacterial activity of a female's exudates, and this subsequently reduces her lifetime reproductive success. We suggest that the costliness of social immunity is a source of evolutionary conflict between breeding adults on a carcass, and that the phoretic communities that the beetles transport between carrion may assist the beetle by offsetting these costs.

Sexual division of antibacterial resource defence in breeding burying beetles, Nicrophorus vespilloides

A key component of parental care involves defending resources destined for offspring from a diverse array of potential interspecific competitors, such as social parasites, fungi and bacteria. 2. Just as with other aspects of parental care, such as offspring provisioning or brood defence, sexual conflict between parents may arise over how to share the costs of this form of care. There has been little previous work, however, to investigate how this particular burden might be shared. 3. Here, we describe a hitherto uncharacterized form of parental care in burying beetles Nicrophorus vespilloides, a species which prepares carrion for its young and faces competition from microbes for this resource. We found that parents defend the carcass with antibacterial anal exudates, and that the antibacterial activity of these exudates is only upregulated following the discovery of a corpse. At the same time, phenoloxidase activity in the anal exudates is downregulated, indicating parallels with the internal insect immune system. 4. In unmanipulated breeding pairs, females had higher antibacterial activity in their anal exudates than males, suggesting sex-specific roles in this aspect of parental care. 5. When we experimentally widowed males, we found that they increased levels of antibacterial activity in their anal exudates. Experimentally widowing females, however, led them to decrease levels of antibacterial activity in their anal exudates. Widowed beetles of each sex thus produced anal exudates of comparable antibacterial activity. We suggest that this flexible division of antibacterial activity may be coordinated by Juvenile Hormone. © 2009 British Ecological Society.

Can the protein costs of bacterial resistance be offset by altered feeding behaviour?

1. Mounting an immune response is likely to be costly in terms of energy and nutrients, and so it is predicted that dietary intake should change in response to infection to offset these costs. The present study focuses on the interactions between a specialist grass-feeding caterpillar species, the African armyworm Spodoptera exempta, and an opportunist bacterium, Bacillus subtilis.
2. The main aims of the study were (i) to establish the macronutrient costs to the insect host of surviving a systemic bacterial infection, (ii) to determine the relative importance of dietary protein and carbohydrate to immune system functions, and (iii) to determine whether there is an adaptive change in the host's normal feeding behaviour in response to bacterial challenge, such that the nutritional costs of resisting infection are offset.
3. We show that the survival of bacterially infected larvae increased with increasing dietary protein-to-carbohydrate (P:C) ratio, suggesting a protein cost associated with bacterial resistance. As dietary protein levels increased, there was an increase in antibacterial activity, phenoloxidase (PO) activity and protein levels in the haemolymph, providing a potential source for this protein cost. However, there was also evidence for a physiological trade-off between antibacterial activity and phenoloxidase activity, as larvae whose antibacterial activity levels were elevated in response to immune activation had reduced PO activity.
4. When given a choice between two diets varying in their P:C ratios, larvae injected with a sub-lethal dose of bacteria increased their protein intake relative to control larvae whilst maintaining similar carbohydrate intake levels. These results are consistent with the notion that S. exempta larvae alter their feeding behaviour in response to bacterial infection in a manner that is likely to enhance the levels of protein available for producing the immune system components and other factors required to resist bacterial infections (‘self-medication’).

Personal immunity versus social immunity

[No abstract available]

Current brood size and residual reproductive value predict offspring desertion in the burying beetle Nicrophorus vespilloides

Life-history theory suggests that offspring desertion can be an adaptive reproductive strategy, in which parents forgo the costly care of an unprofitable current brood to save resources for future reproduction. In the burying beetle, Nicrophorus vespilloides, parents commonly abandon their offspring to the care of others, resulting in female-only care, male-only care, brood parasitism, and the care of offspring sired by satellite males. Furthermore, when there is biparental care, males routinely desert the brood before larval development is complete, leaving females behind to tend their young. We attempted to understand these patterns of offspring desertion by using laboratory experiments to compare the fitness costs associated with parental care for each sex and the residual reproductive value of the 2 sexes. We also tested whether current brood size and residual reproductive value together predicted the incidence of brood desertion. We found that males and females each sustained fecundity costs as a consequence of caring for larvae and that these costs were of comparable magnitude. Nevertheless, males had greater residual reproductive value than females and were more likely than females to desert experimental broods. Our results can explain why males desert the brood earlier than females in nature and why female-only care is more common than male-only care. They also suggest that the tipping point from brood parasitism or satellite male behavior to communal breeding (and vice versa) depends on the value of the current brood relative to residual reproductive value.

Selection for cuticular melanism reveals immune function and life-history trade-offs in Spodoptera littoralis

Several insect species show an increase in cuticular melanism in response to high densities. In some species, there is evidence that this melanism is correlated with an up-regulation of certain immune system components, particularly phenoloxidase (PO) activity, and with the down-regulation of lysozyme activity, suggesting a trade-off between the two traits. As melanism has a genetic component, we selected both melanic and nonmelanic lines of the phase-polyphenic lepidopteran, Spodoptera littoralis, in order to test for a causative genetic link between melanism, PO activity and lysozyme activity, and to establish if there are any life-history costs associated with the melanic response. We found that, in fact, melanic lines had lower PO activity and higher lysozyme activity than nonmelanic lines, confirming a genetic trade-off between the two immune responses, but also indicating a genetic trade-off between melanism and PO activity. In addition, we found that lines with high PO activity had slower development rates suggesting that investment in PO, rather than in melanism, is costly.

Male morph predicts investment in larval immune function in the dung beetle, Onthophagus taurus

Investment in immunity is costly, so that resource-based trade-offs between immunity and sexually selected ornaments might be expected. The amount of resources that an individual can invest in each trait will be limited by the total resources available to them. It would therefore be informative to investigate how investment in immune function changes during growth or production of the sexual trait as resources are diverted to it. Using the dung beetle, Onthophagus taurus, which displays both sexual and male dimorphism in horn size, we examined changes in one measure of immune function, phenoloxidase (PO) activity, in the hemolymph of larvae prior to and during horn growth. We found that PO levels differed between small- and large-horned males throughout the final instar prior to the point where investment in horn growth was taking place. PO levels in females were intermediate to the 2 male morphs. These differences could not be accounted for by differences in condition, measured as hemolymph protein levels and weight. We suggest that the observed differences might be associated with sex- and morph-specific variation in juvenile hormone levels.

Costs of resistance: genetic correlations and potential trade-offs in an insect immune system

Theory predicts that natural selection will erode additive genetic variation in fitness-related traits. However, numerous studies have found considerable heritable variation in traits related to immune function, which should be closely linked to fitness. This could be due to trade-offs maintaining variation in these traits. We used the Egyptian cotton leafworm, Spodoptera littoralis, as a model system to examine the quantitative genetics of insect immune function. We estimated the heritabilities of several different measures of innate immunity and the genetic correlations between these immune traits and a number of life history traits. Our results provide the first evidence for a potential genetic trade-off within the insect immune system, with antibacterial activity (lysozyme-like) exhibiting a significant negative genetic correlation with haemocyte density, which itself is positively genetically correlated with both haemolymph phenoloxidase activity and cuticular melanization. We speculate on a potential trade-off between defence against parasites and predators, mediated by larval colour, and its role in maintaining genetic variation in traits under natural selection.

Data mining approach identifies research priorities and data requirements for resolving the red algal tree of life

Background. The assembly of the tree of life has seen significant progress in recent years but algae and protists have been largely overlooked in this effort. Many groups of algae and protists have ancient roots and it is unclear how much data will be required to resolve their phylogenetic relationships for incorporation in the tree of life. The red algae, a group of primary photosynthetic eukaryotes of more than a billion years old, provide the earliest fossil evidence for eukaryotic multicellularity and sexual reproduction. Despite this evolutionary significance, their phylogenetic relationships are understudied. This study aims to infer a comprehensive red algal tree of life at the family level from a supermatrix containing data mined from GenBank. We aim to locate remaining regions of low support in the topology, evaluate their causes and estimate the amount of data required to resolve them. Results. Phylogenetic analysis of a supermatrix of 14 loci and 98 red algal families yielded the most complete red algal tree of life to date. Visualization of statistical support showed the presence of five poorly supported regions. Causes for low support were identified with statistics about the age of the region, data availability and node density, showing that poor support has different origins in different parts of the tree. Parametric simulation experiments yielded optimistic estimates of how much data will be needed to resolve the poorly supported regions (ca. 103 to ca. 104 nucleotides for the different regions). Nonparametric simulations gave a markedly more pessimistic image, some regions requiring more than 2.8 105 nucleotides or not achieving the desired level of support at all. The discrepancies between parametric and nonparametric simulations are discussed in light of our dataset and known attributes of both approaches. Conclusions. Our study takes the red algae one step closer to meaningful inclusion in the tree of life. In addition to the recovery of stable relationships, the recognition of five regions in need of further study is a significant outcome of this work. Based on our analyses of current availability and future requirements of data, we make clear recommendations for forthcoming research.

Intertidal molluscan and algal species richness around the UK coast

Geographically referenced databases of species records are becoming increasingly available. Doubts over the heterogeneous quality of the underlying data may restrict analyses of such collated databases. We partitioned the spatial variation in species richness of littoral algae and molluscs from the UK National Biodiversity Network database into a smoothed mesoscale component and a local component. Trend surface analysis (TSA) was used to define the mesoscale patterns of species richness, leaving a local residual component that lacked spatial autocorrelation. The analysis was based on 10 km grid squares with 115035 records of littoral algae (729 species) and 66879 records of littoral molluscs (569 species). The TSA identified variation in algal and molluscan species richness with a characteristic length scale of approximately 120 km. Locations of the most species-rich grid squares were consistent with the southern and western bias of species richness in the UK marine flora and fauna. The TSA also identified areas which showed significant changes in the spatial pattern of species richness: breakpoints, which correspond to major headlands along the south coast of England. Patterns of algal and molluscan species richness were broadly congruent. Residual variability was strongly influenced by proxies of collection effort, but local environmental variables including length of the coastline and variability in wave exposure were also important. Relative to the underlying trend, local species richness hotspots occurred on all coasts. While there is some justification for scepticism in analyses of heterogeneous datasets, our results indicate that the analysis of collated datasets can be informative.

Like a rolling stone: the mobility of maerl (Corallinaceae) and the neutrality of the associated assemblages

Beds of nonattached coralline algae (maerl or rhodoliths) are widespread and considered relatively species rich. This habitat is generally found in areas where there is chronic physical disturbance such that maerl thalli are frequently moved. Little is known, however, about how natural disturbance regimes affect the species associated with maerl. This study compared the richness, animal abundance, and algal biomass of maerl-associated species over a two-year period in a wave-disturbed bed and a sheltered maerl bed. Changes in associated species over time were assessed for departures from a neutral model in which the dissimilarity between samples reflects random sampling from a common species pool. Algal biomass and species richness at the wave-exposed site and on stabilized maerl at the sheltered site were reduced at times of higher wind speeds. The changes in species richness were not distinguishable from a neutral model, implying that algal species were added at random to the assemblage as the level of disturbance lessened. Results for animal species were more mixed. Although mobile species were less abundant during windy periods at the exposed site, both neutral and non-neutral patterns were evident in the assemblages. Artificial stabilization of maerl had inconsistent effects on the richness of animals but always resulted in more attached algal species. While the results show that the response of a community to disturbance can be neutral, the domain of neutral changes in communities may be relatively small. Alongside non-neutral responses to natural disturbance, artificial stabilization always resulted in an assemblage that was more distinct than would be expected under random sampling from a common pool. Community responses to stabilization treatments did not consistently follow the predictions of the dynamic equilibrium model, the intermediate disturbance model, or a facilitation model. These inconsistencies may reflect site-specific variation in both the disturbance regime and the adjacent habitats that provide source populations for many of the species found associated with maerl.

Evaluating signatures of glacial refugia for North Atlantic benthic marine taxa

A goal of phylogeography is to relate patterns of genetic differentiation to potential historical geographic isolating events. Quaternary glaciations, particularly the one culminating in the Last Glacial Maximum ~21 ka (thousands of years ago), greatly affected the distributions and population sizes of temperate marine species as their ranges retreated southward to escape ice sheets. Traditional genetic models of glacial refugia and routes of recolonization include these predictions: low genetic diversity in formerly glaciated areas, with a small number of alleles/haplotypes dominating disproportionately large areas, and high diversity including "private" alleles in glacial refugia. In the Northern Hemisphere, low diversity in the north and high diversity in the south are expected. This simple model does not account for the possibility of populations surviving in relatively small northern periglacial refugia. If these periglacial populations experienced extreme bottlenecks, they could have the low genetic diversity expected in recolonized areas with no refugia, but should have more endemic diversity (private alleles) than recently recolonized areas. This review examines evidence of putative glacial refugia for eight benthic marine taxa in the temperate North Atlantic. All data sets were reanalyzed to allow direct comparisons between geographic patterns of genetic diversity and distribution of particular clades and haplotypes including private alleles. We contend that for marine organisms the genetic signatures of northern periglacial and southern refugia can be distinguished from one another. There is evidence for several periglacial refugia in northern latitudes, giving credence to recent climatic reconstructions with less extensive glaciation.