Phenotypic shifts caused by predation: selection or life-history shifts?

Predators can impose both selection and life-history shifts in prey populations. Because both processes may affect phenotypic distributions, the estimates of selection differentials may be biased. We carried out two field experiments to disentangle these separate effects. We studied whether dragonfly predation by Aeshna cyanea changes the distributions in body size and lamellae morphology in the damselfly Lestes sponsa. Damselflies have caudal lamellae which are used in escapes by swimming. In a first experiment, we manipulated predator presence (No Aeshna, Encaged Aeshna or Free-ranging Aeshna) and stopped the experiment when all larvae had moulted once. In a second experiment, larvae were confronted with a Free-ranging Aeshna but collected before moulting, and survivors were compared with a control sample taken at the start of the experiment. The presence of Aeshna largely reduced the survival probabilities of the Lestes larvae at a very similar rate in both experiments. Daily survival probabilities did not differ between the No Aeshna and Encaged Aeshna treatments. In the Free-ranging Aeshna treatment of the first experiment, size was reduced compared to the other two treatments, creating a significant apparent selection differential. This was probably mainly due to predator-induced reduced growth because in the second experiment, where growth effects were excluded, size of the survivors did not differ from the control sample. In both experiments there was a significant selection pressure for larger lamellae. Standardized directional selection differentials were similar in both experiments (0.57 and 0.28 phenotypic standard deviation units). No survival selection on lamellae shape was detected. These results are in agreement with previous findings that lamellae size, but not lamellae shape, enhances swimming performance and thereby predator escape in this species. This revised version was published online in July 2006 with corrections to the Cover Date.     

Dimorphic male midshipman fish: reduced sexual selection or sexual selection for reduced characters?

In most taxa with male dimorphisms, some males are large inbody size with exaggerated secondary sexual characters (exaggeratedmorph), whereas other males in the same population are smalland have reduced secondary sexual characters (reduced morph).What selective pressures cause male dimorphisms? Reduced morphologiesmay result when a) some males develop a morphology that, inthe absence of sexual selection pressures for an exaggeratedmorphology, reduces energetic and developmental costs and/orb) some males opt for an alternative morphology that does wellat an alternative behavioral tactic such as cuckoldry. The 2mechanisms could act together, but each alone is theoreticallysufficient to drive dimorphisms. Here, we tested hypothesis"b" (sexual selection for reduced characters) in the plainfinmidshipman fish, Porichthys notatus. Behavioral plasticity betweenterritoriality and cuckoldry in an exaggerated male morph (typeI) allows for a direct comparison of cuckoldry by exaggeratedmorph males to cuckoldry by reduced morph (type II) males. Comparedwith type I cuckolders, type II cuckolders were able to remainnear the nest for longer periods before being chased by theterritorial type I male, suggesting that the reduced type IImorphology allows type II males to prolong the time before attackby territorial males. Combined with other studies showing arole of sexual selection in maintaining the exaggerated morph,the data support the "sexual selection for reduced characters"hypothesis and elucidate how sexual selection can act in differentways on different males to maintain 2 male morphologies withina single species.     

Avoidance of oxidative-stress perturbation in yeast bioprocesses by proteomic and genomic biostrategies?

AIMS: Bioprocess oxidative stress caused by many reactive oxygen species (ROS) can lead to largely irreversible perturbation of yeast bioprocesses. These include the production of proteins derived from recombinant DNA yeast technology (aerobically grown Saccharomyces cerevisiae). These proteins include rennin, amyloglucosidases (glucamylases), interferons, interleukins, insulin, monoclonal antibodies, tissue plasminogen activators (t-PA), sexually transmitted disease antigens, and measles, mumps and rubella antigens, growth hormones, somatotropin, blood clotting factors VIII and XIII. In addition, there may be a demand for severe acute respiratory syndrome-coronavirus antigens, hepatitis A, B and C viral-selected antigens, HIV retroviral antigens, influenza antigens, trypanosomal antigens, and foot and mouth disease antigens. Prevention of oxidative stress has been achieved by application of antioxidant redox metalloenzymes such as superoxide dismutases (containing Cu/Zn cytosolic, Mn mitochondrial and Fe bacterial) glutathione peroxidases (and other Se-containing proteins and enzymes such as the thioredoxins), catalases (Fe-containing), cytochrome c peroxidases (Fe-containing), ceruloplasmins (Cu-containing), metallothionines (these cysteine thiol-rich proteins bind ions of cadmium and mercury) and tyrosinases(Cu-containing). METHODS AND RESULTS: ROS are generated inadvertently by single metal valency couples such as FeII/FeIII and by FeIII/FeV present in 2700 (including 57 human) isoforms in cytochromes P450 mixed-function oxidases (EC 1.14.14.1; O2 : mono-oxygenase NADPH/NADH requiring). In addition, mixed-metal couples such as valency unmatched forms in CuI/FeII and FeIII/MnIV can recycle electrons. Moreover, proteins/protein chaperone couples can recycle electrons, often where futile-recycling systems have been instigated. Furthermore, oxidized membrane phospholipids (R) can form ROOH (lipid hydroperoxides) and ROH (lipid alkoxides) that can generate ROS through Fenton chemistry (iron-catalysed) chain reactions. Utilization of chain-breaking antioxidants such as vitamin E (alpha-tocopherol) in the lipid phase and vitamin C (ascorbate) in the aqueous phase can terminate these ROS-producing reactions. CONCLUSIONS: The main significance of the study is that proteomic strategies of relief from bioprocess perturbation by ROS of yeast fermentations (used to manufacture proteins required in the food and therapeutic bioindustries) may become possible through addition of selected proteins (including metalloenzymes). The main impact of the study is that the utilization of genetically modified (GM) yeast produced by recombinant DNA technology genomic strategies could circumvent the bioprocessing problems that otherwise result from the bioprocess perturbations: this is as a result of oxidative stress caused by ROS, which is avoidable by deployment of appropriate antioxidants such as vitamins E, C and D (and antioxidant proteins and enzymes often of microbial origin via recombinant DNA technology).     

What Happens after Inbreeding Avoidance? Inbreeding by Rejected Relatives and the Inclusive Fitness Benefit of Inbreeding Avoidance

Avoiding inbreeding, and therefore avoiding inbreeding depression in offspring fitness, is widely assumed to be adaptive in systems with biparental reproduction. However, inbreeding can also confer an inclusive fitness benefit stemming from increased relatedness between parents and inbred offspring. Whether or not inbreeding or avoiding inbreeding is adaptive therefore depends on a balance between inbreeding depression and increased parent-offspring relatedness. Existing models of biparental inbreeding predict threshold values of inbreeding depression above which males and females should avoid inbreeding, and predict sexual conflict over inbreeding because these thresholds diverge. However, these models implicitly assume that if a focal individual avoids inbreeding, then both it and its rejected relative will subsequently outbreed. We show that relaxing this assumption of reciprocal outbreeding, and the assumption that focal individuals are themselves outbred, can substantially alter the predicted thresholds for inbreeding avoidance for focal males. Specifically, the magnitude of inbreeding depression below which inbreeding increases a focal male’s inclusive fitness increases with increasing depression in the offspring of a focal female and her alternative mate, and it decreases with increasing relatedness between a focal male and a focal female’s alternative mate, thereby altering the predicted zone of sexual conflict. Furthermore, a focal male’s inclusive fitness gain from avoiding inbreeding is reduced by indirect opportunity costs if his rejected relative breeds with another relative of his. By demonstrating that variation in relatedness and inbreeding can affect intra- and inter-sexual conflict over inbreeding, our models lead to novel predictions for family dynamics. Specifically, parent-offspring conflict over inbreeding might depend on the alternative mates of rejected relatives, and male-male competition over inbreeding might lead to mixed inbreeding strategies. Making testable quantitative predictions regarding inbreeding strategies occurring in nature will therefore require new models that explicitly capture variation in relatedness and inbreeding among interacting population members.     

Are the preference and selection patterns of hermit crabs for gastropod shells species- or site-specific?

Experimental analyses of hermit crabs and their preferences for shells are essential to understand the intrinsic relationship of the crabs' dependence on shells, and may be useful to explain their shell use pattern in nature. The aim of this study was to evaluate the effect of crab species and site on the pattern of shell use, selection, and preference in the south-western Atlantic hermit crabs Pagurus brevidactylus and Pagurus criniticornis, comparing sympatric and allopatric populations. Differently from the traditional approach to evaluate shell preference by simply determining the shell selection pattern (i.e., the number of shells of each type selected), preference was defined (according to [Liszka, D., Underwood, A.J., 1990. An experimental design to determine preferences for gastropod shells by a hermit-crab. J. Exp. Mar. Biol. Ecol., 137(1), 47–62]) by the comparison of the number of crabs changing for a particular shell type when three options were given (Cerithium atratum, Morula nodulosa, and Tegula viridula) with the number of crabs changing for this same type when only this type was offered. The effect of crab species was tested at Cabelo Gordo Beach, where P. brevidactylus was found occupying shells of C. atratum, M. nodulosa, and T. viridula in similar frequencies, whereas P. criniticornis occupied predominantly shells of C. atratum. In laboratory experiments the selection patterns of the two hermit-crab species for these three gastropods were different, with P. criniticornis selecting mainly shells of C. atratum, and P. brevidactylus selecting more shells of M. nodulosa. The shell preference was also dependent on crab species, with P. criniticornis showing a clear preference for shells of C. atratum, whereas P. brevidactylus did not show a preference for any of the tested shells. The effect of site was tested for the two species comparing data from Cabelo Gordo to Preta (P. brevidactylus) and Araçá beaches (P. criniticornis). The pattern of shell use, selection, and preference was demonstrated to be dependent on site only for P. brevidactylus. The results also showed that the shell use pattern of P. criniticornis can be explained by its preference at both sites, whereas for P. brevidactylus it occurred only at Cabelo Gordo, where the absence of preference was correlated with the similar use of the three gastropod species studied. Finally, the results showed that the shell selection pattern cannot be considered as a measure of shell preference, since it overestimates crab selectivity.     

Paradoxes of tumour complexity: somatic selection,vulnerability by design,or infectious aetiology?

The aetiology of cancer involves intricate cellular and molecular mechanisms that apparently emerge on the short timescale of a single lifetime. Some of these traits are remarkable not only for their complexity, but also because it is hard to conceive selection pressures that would favour their evolution within the local competitive microenvironment of the tumour. Examples include ‘niche construction’ (re‐programming of tumour‐specific target sites) to create permissive conditions for distant metastases; long‐range feedback loops of tumour growth; and remarkably ‘plastic’ phenotypes (e.g. density‐dependent dispersal) associated with metastatic cancer. These traits, which we term ‘paradoxical tumour traits’, facilitate the long‐range spread or long‐term persistence of the tumours, but offer no apparent benefit, and might even incur costs in the competition of clones within the tumour. We discuss three possible scenarios for the origin of these characters: somatic selection driven by specific selection regimes; non‐adaptive emergence due to inherent vulnerabilities in the organism; and manipulation by putative transmissible agents that contribute to and benefit from these traits. Our work highlights a lack of understanding of some aspects of tumour development, and offers alternative hypotheses that might guide further research.     

Tracking prey or tracking the prey's resource? Mechanisms of movement and optimal habitat selection by predators

We synthesize previous theory on ideal free habitat selection to develop a model of predator movement mechanisms, when both predators and prey are mobile. We consider a continuous environment with an arbitrary distribution of resources, randomly diffusing prey that consume the resources, and predators that consume the prey. Our model introduces a very general class of movement rules in which the overall direction of a predator's movement is determined by a variable combination of (i) random diffusion, (ii) movement in the direction of higher prey density, and/or (iii) movement in the direction of higher density of the prey's resource. With this model, we apply an adaptive dynamics approach to two main questions. First, can it be adaptive for predators to base their movement solely on the density of the prey's resource (which the predators do not consume)? Second, should predator movements be exclusively biased toward higher densities of prey/resources, or is there an optimal balance between random and biased movements? We find that, for some resource distributions, predators that track the gradient of the prey's resource have an advantage compared to predators that track the gradient of prey directly. Additionally, we show that matching (consumers distributed in proportion to resources), overmatching (consumers strongly aggregated in areas of high resource density), and undermatching (consumers distributed more uniformly than resources) distributions can all be explained by the same general habitat selection mechanism. Our results provide important groundwork for future investigations of predator-prey dynamics.     

Avoidance of willows from moderately polluted area by leaf beetle,Melasoma lapponica: effects of emission or induced resistance?

We studied preference for willows along a pollution gradient on the Kola Peninsula, Russia, by the leaf beetle, Melasoma lapponica. Multiple tests with leaf disks demonstrated low preference for Salix borealis, S. caprea and S. phylicifolia from the plot situated 14 km from the smelter, in comparison with conspecific plants from plots situated at 1 and 29 km distances. This pattern was observed when testing beetles orginating from any plot both in 1993 and 1994, using both young and mature leaves of S. borealis. Although fumigation of S. borealis with realistic SO₂ concentration (100 g/m³) increased plant palatability, preference for plants from our study plots did not correlate with plot-specific mean SO₂ concentrations. Furthermore, no correlation with foliar concentrations of the main metal pollutants (Ni and Cu) was found. Palatability of plants was negatively correlated with population density of M. lapponica, which peaked in the moderately polluted plot 14 km from the smelter. Within this plot, beetles clearly preferred non-damaged bushes of S. borealis to previously damaged bushes. We therefore conclude that low preference of S. borealis from the moderately polluted area was caused by plant resistance induced by severe damage from M. lapponica in previous years rather than by pollution impact. However, S. caprea and S. phylicifolia had little damage from M. lapponica, and low palatability of these species in the moderately polluted plot suggests changes in plant quality similar to changes in heavily damaged bushes of S. borealis.     

Amyloidogenic domains, prions and structural inheritance: rudiments of early life or recent acquisition?

Amyloids are self-assembled fibre-like beta-rich protein aggregates. Amyloidogenic prion proteins propagate amyloid state in vivo and transmit it via infection or in cell divisions. While amyloid aggregation may occur in the absence of any other proteins, in vivo propagation of the amyloid state requires chaperone helpers. Yeast prion proteins contain prion domains which include distinct aggregation and propagation elements, responsible for these functions. Known aggregation and propagation elements are short in length and composed of relatively simple sequences, indicating possible ancient origin. Prion-like self-assembled structures could be involved in the initial steps of biological compartmentalization in early life.     

Is flight architecture determined by physical constraints or by natural selection?: the case of the midge Chironomus plumosus

Males of the midge Chironomus plumosus fly solely to mate. They maintain station for long periods in moving air. Females patrol in search of males and, after receiving a spermatophore, fly to oviposition sites. The requirements of flight in males and females are therefore fundamentally different. Females are larger than males, on average, so these differences could stem from scaling rules governing the geometry of space. The same explanation might apply to flight differences within the sexes and even to peculiarities of flight architecture in C. plumosus compared to other flying animals. In other words, might flight design be accounted for entirely by mechanical constraints without recourse to natural selection?
To test this hypothesis, the power output of C. plumosu was measured as size-specific muscle mass. Contrary to expectation, little evidence was found of scaling effects in this measure of power. Despite its being among the smallest of animals to fly, C. plumosus turns out to have the largest mass of flight muscle, relative to body size, yet found among animals. Differences both between C. plumosus and other species and within C. plumosus are, in general, more readily accounted for by the requirements of the mating system within the viscous universe encountered by small flying animals. We conclude that it is natural selection rather than mechanical constraint that is the primary influence determining the architecture of flight in this small animal.     

Cereal phytochromes: targets of selection, targets for manipulation?

Plants respond to shading through an adaptive syndrome termed shade avoidance. In high-density crop plantings, shade avoidance generally increases extension growth at the expense of yield and can be at odds with the agronomic performance of the crop as a whole. Studies in Arabidopsis are beginning to reveal the essential role phytochromes play in regulating this process and to identify genes underlying the response. In this article, we focus on how phytochrome signaling networks have been targeted in cereal breeding programs in the past and discuss the potential to alter these pathways through breeding and transgenic manipulation to develop crops that perform better under typical high density conditions.     

Manipulation of host behaviour by parasites: a weakening paradigm?

New scientific paradigms often generate an early wave of enthusiasm among researchers and a barrage of studies seeking to validate or refute the newly proposed idea. All else being equal, the strength and direction of the empirical evidence being published should not change over time, allowing one to assess the generality of the paradigm based on the gradual accumulation of evidence. Here, I examine the relationship between the magnitude of published quantitative estimates of parasite-induced changes in host behaviour and year of publication from the time the adaptive host manipulation hypothesis was first proposed. Two independent data sets were used, both originally gathered for other purposes. First, across 137 comparisons between the behaviour of infected and uninfected hosts, the estimated relative influence of parasites correlated negatively with year of publication. This effect was contingent upon the transmission mode of the parasites studied. The negative relationship was very strong among studies of parasites which benefit from host manipulation (transmission to the next host occurs by predation on an infected intermediate host), i.e. among studies which were explicit tests of the adaptive manipulation hypothesis. There was no correlation with year of publication among studies on other types of parasites which do not seem to receive benefits from host manipulation. Second, among 14 estimates of the relative, parasite-mediated increase in transmission rate (i.e. increases in predation rates by definitive hosts on intermediate hosts), the estimated influence of parasites again correlated negatively with year of publication. These results have several possible explanations, but tend to suggest biases with regard to what results are published through time as accepted paradigms changed.     

Regulation of aphid populations by aphidiid wasps: does parasitoid foraging behaviour or hyperparasitism limit impact?

Aphidiid parasitoids (Hymenoptera: Aphidiidae) of aphids generally exploit only a small percentage of the available host resources in the field. This limited impact on aphid populations has often been explained as a consequence of hyperparasitism. We propose that a wasp's reproductive strategy, as opposed to hyperparasitism, is the dominant factor in aphidiid population dynamics. A wasp's foraging efficiency and oviposition decisions are influenced by several variables, including searching behaviour between and within patches, host choice (as modified by the aphids' defensive behaviours), and plant structural complexity. Two broadly different patterns of host exploitation have evolved in aphidiid wasps in relation to ant-aphid mutualism. Firstly, in species that are exposed to predation and hyperparasitism, a female may leave a patch before all suitable hosts are parasitized. Because predators and hyperparasitoids tend to aggregate at high aphid or aphidiid densities, or in response to aphid honeydew, this strategy enables females to reduce offspring mortality by spreading the risk over several host patches. Secondly, in species that have evolved mechanisms to avoid aggression by mutualistic ants, females are able to exploit a hyperparasitoid-free resource space. Such species may concentrate their eggs in only a few aphid colonies, which are thus heavily exploited. Although hyperparasitism of species in the first group tends to reach high levels, its overall impact on aphid-aphidiid population dynamics is probably limited by the low average fecundity of most hyperparasitoids. We discuss the foraging patterns of aphidiid wasps in relation to aphid population regulation in general, and to classical biological control in particular. We argue that a parasitoid's potential to regulate the host population is largely determined by its foraging strategy. In an exotic parasitoid, a behavioural syndrome that has evolved and presumably is adaptive in a more diverse (native) environment may, in a more uniform (managed) environment, result in suboptimal patch-leaving and oviposition decisions, and possibly increased resource usage.     

Isolated remnant or recent introduction? Estimating the provenance of Yellingbo Leadbeater's possums by genetic analysis and bottleneck simulation

Effective conservation management requires that genetically divergent populations potentially harbouring important local adaptations be identified and maintained as separate management units. In the case of the endangered Australian Leadbeater's possum (Gymnobelideus leadbeateri), an arboreal marsupial endemic to Victoria, uncertainty over the evolutionary origin of a potentially important extant wild population recently discovered in atypical habitat (lowland swamp) at Yellingbo is hampering such efforts. The population is rumoured to be a recent introduction. Microsatellite allele frequencies at Yellingbo differed substantially from those in sampled populations in montane ash forest (F_ST between 0.23 and 0.36), and Bayesian clustering analyses of genotypes strongly separated them (K = 2). We conducted a suite of bottlenecking tests which all indicated that Yellingbo had undergone a recent reduction in size. The extent to which the distinctiveness of Yellingbo animals might be expected solely through bottlenecking associated with a recent introduction, was tested by simulating population–history scenarios seeded with genotypes from candidate wild and captive sources. No bottleneck scenario reproduced anything approaching the genetic distinction of the Yellingbo population, with all structure analyses placing Yellingbo in a separate cluster to simulated populations (K = 2, minimum F_ST = 0.13). These results suggest that Yellingbo does not share recent ancestry with other extant populations and instead may be a remnant of an otherwise extinct gene pool. Importantly, this may include genes involved in adaptation to a lowland swamp environment, substantially adding to the conservation importance of this population, and suggesting that separate management may be prudent until evidence suggests otherwise.     

Fig choice by the pollinator of a gynodioecious fig: selection to rush,or intersexual mimicry?

Due to selection to maximise the benefits accrued, conflicts of interest exist between mutualists. An extreme conflict occurs between gynodioecious fig trees and their pollinating wasps. Female wasps, which are required for pollination, cannot reproduce in female figs and should be selected to avoid them. We investigate fig choice in Liporrhopalum tentacularis , pollinator of Ficus montana . We show that entry rates are independent of fig sex and diameter, with wasps instead entering the first fig encountered (although significant between-tree differences in entry rates to figs were observed). This suggests that wasps are unable to discriminate between the sexes because of selection for inter-sexual fig mimicry. In conjuction with data on other species pairs, these findings imply that that the resolution of the conflict is ultimately under the tree's control, with the level of mimicry evolving in response to both the propensity of wasps to discriminate and the costs of them doing so.