Gender-specific differences in cannibalism between a laboratory strain and a field strain of a predatory mite.

Many phytoseiid species, including Phytoseiulus persimilis, are known to engage in cannibalism when food is scarce and when there is no possibility to disperse. In nature adult females of P. persimilis are known to disperse when prey is locally depleted. Males, in contrast, are expected to stay and wait for potential mates to mature. During this phase, males can obtain food by cannibalizing. Therefore, we hypothesize that male P. persimilis exhibit a higher tendency to cannibalize than females. Because rearing conditions in the laboratory usually prevent dispersal, prolonged culturing may also affect cannibalistic behavior. We hypothesize that this should especially affect cannibalism by females, because they consume far more food. We tested these hypotheses by comparing males and females from two strains, one of which had been in culture for over 20 years, whereas the other was recently collected from the field. It is known that this predator can discriminate between kin and non-kin and prefers cannibalizing the latter, hence to construct lines with high relatedness we created isofemale lines of these two original strains. We subsequently tested to what extent the adult females and males of the original strains and the isofemale lines cannibalized conspecific larvae from the same strain/line in a closed system. Relatedness with the victims did not affect cannibalistic behavior, but males engaged more often in cannibalism than females, and females of the laboratory strain engaged more in cannibalism than those of the field strain, both in agreement with our ideas. We hypothesize that the difference in cannibalism between the two genders will increase when they have the alternative to disperse.

Detection of genetic incompatibilities in non-model systems using simple genetic markers: hybrid breakdown in the haplodiploid spider mite Tetranychus evansi.

When two related species interbreed, their hybrid offspring frequently suffer from reduced fitness. The genetics of hybrid incompatibility are described by the Bateson-Dobzhansky-Muller (BDM) model, where fitness is reduced by epistatic interactions between alleles of heterospecific origin. Unfortunately, most empirical evidence for the BDM model comes from a few well-studied model organisms, restricting our genetic understanding of hybrid incompatibilities to limited taxa. These systems are predominantly diploid and incompatibility is often complete, which complicates the detection of recessive allelic interactions and excludes the possibility to study viable or intermediate stages. Here, we advocate research into non-model organisms with haploid or haplodiploid reproductive systems and incomplete hybrid incompatibility because (1) dominance is absent in haploids and (2) incomplete incompatibility allows comparing affected with unaffected individuals. We describe a novel two-locus statistic specifying the frequency of individuals for which two alleles co-occur. This approach to studying BDM incompatibilities requires genotypic characterization of hybrid individuals, but not genetic mapping or genome sequencing. To illustrate our approach, we investigated genetic causes for hybrid incompatibility between differentiated lineages of the haplodiploid spider mite Tetranychus evansi, and show that strong, but incomplete, hybrid breakdown occurs. In addition, by comparing the genotypes of viable hybrid males and inviable hybrid male eggs for eight microsatellite loci, we show that nuclear and cytonuclear BDM interactions constitute the basis of hybrid incompatibility in this species. Our approach opens up possibilities to study BDM interactions in non-model taxa, and may give further insight into the genetic mechanisms behind hybrid incompatibility.

Context-dependent alarm signalling in an insect.

Animals often respond to danger by raising alarm to inform others. Alarm signals come in many different forms, such as visual or mechanical display, sound or odour. Some animals produce vocal alarm signals that vary with the level of danger. For chemical alarm signals, virtually nothing is known about such context-dependent signalling due to a general notion that alarm pheromones have fixed compositions. Here, we show that larvae of the Western Flower Thrips (Frankliniella occidentalis) produce an alarm pheromone whose composition varies with the level of danger they face: the presence of a relatively harmless predator or a very dangerous predator, that is either actually attacking or not. The frequency of alarm pheromone excretion increases with the level of danger. Moreover, the composition of excreted alarm pheromone varies in the relationship between total and relative amount of the putative two components, decyl acetate (DAc) and dodecyl acetate (DDAc). When pheromone is excreted with a predator present but not attacking, the percentage DDAc increases with the total amount of pheromone. When a predator does attack, however, the relationship between percentage DDAc and total amount of pheromone is reversed. Taken together, the alarm signal of thrips larvae appears to be context dependent, which to our knowledge is the first report of context-dependent composition of an alarm pheromone.

Inbreeding depression and purging in a haplodiploid: gender-related effects.

Compared with diploid species, haplodiploids suffer less inbreeding depression because male haploidy imposes purifying selection on recessive deleterious alleles. However, alleles of genes only expressed in the diploid females are protected in heterozygous individuals. This leads to the prediction that haplodiploids suffer more from inbreeding effects on life-history traits controlled by genes with female-limited expression. To test this, we used a wild population of the haplodiploid mite Tetranychus urticae. First, negative effects of inbreeding were investigated by comparing maturation rate, juvenile survival, oviposition rate and longevity between lines created by three generations of either outbreeding or mother-son inbreeding. Second, purging through inbreeding was investigated by comparing the intensity of inbreeding depression between outbred families with known inbreeding/outbreeding mating histories. Negative effects of inbreeding and evidence for purging were found for the female trait oviposition rate, but not for juvenile survival and longevity. Both male and female maturation rate were negatively affected by inbreeding, most likely due to maternal effects because inbred offspring of outbred mothers was not affected. These results support the hypothesis that, in haplodiploids inbreeding effects and genetic variation due to deleterious recessive alleles may depend on gender.

Mechanisms and ecological consequences of plant defence induction and suppression in herbivore communities.

BACKGROUND: Plants are hotbeds for parasites such as arthropod herbivores, which acquire nutrients and energy from their hosts in order to grow and reproduce. Hence plants are selected to evolve resistance, which in turn selects for herbivores that can cope with this resistance. To preserve their fitness when attacked by herbivores, plants can employ complex strategies that include reallocation of resources and the production of defensive metabolites and structures. Plant defences can be either prefabricated or be produced only upon attack. Those that are ready-made are referred to as constitutive defences. Some constitutive defences are operational at any time while others require activation. Defences produced only when herbivores are present are referred to as induced defences. These can be established via de novo biosynthesis of defensive substances or via modifications of prefabricated substances and consequently these are active only when needed. Inducibility of defence may serve to save energy and to prevent self-intoxication but also implies that there is a delay in these defences becoming operational. Induced defences can be characterized by alterations in plant morphology and molecular chemistry and are associated with a decrease in herbivore performance. These alterations are set in motion by signals generated by herbivores. Finally, a subset of induced metabolites are released into the air as volatiles and function as a beacon for foraging natural enemies searching for prey, and this is referred to as induced indirect defence. SCOPE: The objective of this review is to evaluate (1) which strategies plants have evolved to cope with herbivores and (2) which traits herbivores have evolved that enable them to counter these defences. The primary focus is on the induction and suppression of plant defences and the review outlines how the palette of traits that determine induction/suppression of, and resistance/susceptibility of herbivores to, plant defences can give rise to exploitative competition and facilitation within ecological communities "inhabiting" a plant. CONCLUSIONS: Herbivores have evolved diverse strategies, which are not mutually exclusive, to decrease the negative effects of plant defences in order to maximize the conversion of plant material into offspring. Numerous adaptations have been found in herbivores, enabling them to dismantle or bypass defensive barriers, to avoid tissues with relatively high levels of defensive chemicals or to metabolize these chemicals once ingested. In addition, some herbivores interfere with the onset or completion of induced plant defences, resulting in the plant's resistance being partly or fully suppressed. The ability to suppress induced plant defences appears to occur across plant parasites from different kingdoms, including herbivorous arthropods, and there is remarkable diversity in suppression mechanisms. Suppression may strongly affect the structure of the food web, because the ability to suppress the activation of defences of a communal host may facilitate competitors, whereas the ability of a herbivore to cope with activated plant defences will not. Further characterization of the mechanisms and traits that give rise to suppression of plant defences will enable us to determine their role in shaping direct and indirect interactions in food webs and the extent to which these determine the coexistence and persistence of species.

Ontogenetic analysis of behavior in the tail suspension test: temporal differences in the emergence of within- and between-session habituation in Swiss mice.

Habituation is an important tool in the investigation of learning/memory throughout life. Despite that, few studies describe habituation from an ontogenetic perspective. Considering that, as soon as they are born, rodents can twist their bodies when lifted by their tails in an attempt to escape, this behavior should be well suited to study habituation behavior from birth to adulthood. Here, we implement a tail suspension test to study the ontogenetic development of habituation in Swiss mice. Our data indicate that a continuous within-session decrease in trunk movements can be observed from postnatal day (P) 10 onwards and that between-sessions habituation (from one day to another) can be observed from P16 onwards. Furthermore, we show that the adult pattern of within- and between-sessions reductions in activity is already present by the beginning of adolescence, at P28. Our results indicate that between-sessions habituation involves a more complex mechanism of memory and learning than within-session habituation, requiring a longer period of brain maturation before it can be displayed.

Environmental effects on the detection of adaptation.

Detecting adaptation involves comparing the performance of populations evolving in different environments. This detection may be confounded by effects due to the environment experienced by organisms prior to the test. We tested whether such confounding effects occur, using spider-mite selection lines on two novel hosts and one ancestral host, after 15 generations of selection. Mites were either sampled directly from the selection lines or subjected to a common juvenile or to a common maternal environment, mimicking the most frequent environmental manipulations. These environments strongly affected all life-history traits. Moreover, the detection of adaptation and correlated responses on the ancestral host was inconsistent among environments in almost 20% of the cases. Indeed, we did not detect responses unambiguously for any life-history trait. This inconsistency was due to differential environmental effects on lines from different selection regimes. Therefore, the detection of adaptation requires a careful control of these environmental effects.

Innate responses of the predatory mite Phytoseiulus persimilis to a herbivore-induced plant volatile.

The responses of the predatory mite P. persimilis to herbivore-induced plant volatiles are at least partly genetically determined. Thus, there is potential for the evolution of this behaviour by natural selection. We tested whether distinct predator genotypes with contrasting responses to a specific herbivore-induced plant volatile, i.e. methyl salicylate (MeSa), could be found in a base population collected in the field (Sicily). To this end, we imposed purifying selection on individuals within iso-female lines of P. persimilis such that the lines were propagated only via the individual that showed either a preference or avoidance of MeSa. The responses of the lines were characterized as the mean proportion of individuals choosing MeSa when given a choice between MeSa and clean air. Significant variation in predator responses was detected among iso-female lines, thus confirming the presence of a genetic component for this behaviour. Nevertheless, we did not find a significant difference in the response to MeSa between the lines that were selected to avoid MeSa and the lines selected to prefer MeSa. Instead, in the course of selection the lines selected to avoid MeSa shifted their mean response towards a preference for MeSa. An inverse, albeit weaker, shift was detected for the lines selected to prefer MeSa. We discuss the factors that may have caused the apparent lack of a response to selection within iso-female line in this study and propose experimental approaches that address them.

Ambulatory dispersal in Tetranychus urticae: an artificial selection experiment on propensity to disperse yields no response.

Dispersal to new hosts is an important process for an invasive herbivore, such as the two-spotted spider mite. A recent study, using artificial selection experiments, has suggested that genetic variation and genetic trade-offs are present for propensity to disperse in this species. However, due to the experimental setup alternative explanations for the response to selection could not be ruled out. Using an altered setup, we investigated whether the propensity for ambulatory dispersal differs genetically between individuals and whether genetic correlations with life-history traits exist. Upward and downward selection on propensity to leave the colony was performed for seven generations in four replicate artificial selection experiments and the results were compared to control lines. No consistent responses to selection were found and no significant effect on life-history traits (oviposition rate, juvenile survival, development rate and number of adult offspring) or sex ratio was present across the replicates. The data suggest that our base population of spider mites harbours at best a low amount of additive genetic variation for this behaviour.

Mate choice promotes inbreeding avoidance in the two-spotted spider mite.

Since inbreeding in Tetranychus urticae can reduce offspring fitness, sexual selection may favour disassortative mate choice with respect to relatedness of the mating partners. We tested whether T. urticae shows this preference for mating with unrelated partners. We chose an experimental set-up with high potential for female choosiness, since females only mate once and are therefore expected to be the choosier gender. An adult virgin female was placed together with two adult males from the same population. One male was unrelated and the other male was related-a brother with whom she had grown up. Significantly more copulations (64%) took place with the unrelated male. Time to mating did not depend on the female-to-male relatedness. The remaining (non-copulating) male tried to interfere with the ongoing mating in the majority of cases, but this interference did not depend on the female-to-male relatedness. These results imply that T. urticae (a) can recognize kin (via genetic and/or environmental similarity) and (b) has the potential to avoid inbreeding through mate choice.

Measurement of Active Motor Threshold Using a Dynamometer During Navigated Transcranial Magnetic Stimulation in a Patient with Postoperative Brain Tumor: Technical Note.

BACKGROUND: Navigated transcranial magnetic stimulation (nTMS) is being used for different purposes in patients with brain tumors. However, the procedure requires a positive electrophysiological response. For patients with negative response in rest conditions, active motor threshold (AMT) may be used. However, sometimes it is difficult to obtain AMT measures owing to inability of the patient to sustain steady muscle contraction. Herein, we describe a simple method by using a hand dynamometer to obtain AMT measures during nTMS session. CASE DESCRIPTION: A woman aged 68 years underwent total removal of a right frontal lobe oligodendroglioma World Health Organization grade II 15 years ago. Cranial magnetic resonance imaging during follow-up revealed local recurrence. In the postoperative period, she developed left upper limb paresis. A postoperative nTMS session was performed for motor electrophysiological evaluation. However, using the standard technique for AMT measurement, the patient was unable to perform sustained muscle contraction as required. A hand dynamometer was used. It allowed sustained muscle contraction for AMT measurement. A counter force for the index finger flexion, the hand support to stabilize hand joints, and a numerical screen serving for both the examiner and the patient as a feedback parameter may explain the success obtained with this simple device. CONCLUSIONS: Although more studies are necessary to validate the method, the hand dynamometer should be considered for patients unable to sustain muscle contraction during AMT measurement.

Judgment of line orientation depends on gender, education, and type of error.

Visuospatial tasks are particularly proficient at eliciting gender differences during neuropsychological performance. Here we tested the hypothesis that gender and education are related to different types of visuospatial errors on a task of line orientation that allowed the independent scoring of correct responses ("hits", or H) and one type of incorrect responses ("commission errors", or CE). We studied 343 volunteers of roughly comparable ages and with different levels of education. Education and gender were significantly associated with H scores, which were higher in men and in the groups with higher education. In contrast, the differences between men and women on CE depended on education. We concluded that (I) the ability to find the correct responses differs from the ability to avoid the wrong responses amidst an array of possible alternatives, and that (II) education interacts with gender to promote a stable performance on CE earlier in men than in women.

Behavioral adjustment to asymmetric reward availability among children with and without ADHD: effects of past and current reinforcement contingencies.

Altered reinforcement sensitivity is hypothesized to underlie symptoms of attention deficit hyperactivity disorder (ADHD). Here we evaluate the behavioral sensitivity of Brazilian children with and without ADHD to a change in reward availability. Forty typically developing children and 32 diagnosed with DSM-IV ADHD completed a signal-detection task in which correct discriminations between two stimuli were associated with different frequencies of reinforcement. The response alternative associated with the higher rate of reinforcement switched, without warning, after 30 rewards were delivered. The task continued until another 30 rewards were delivered. Both groups of children developed a response bias toward the initially more frequently reinforced alternative. This effect was larger in the control group. The response allocation of the two groups changed following the shift in reward availability. Over time the ADHD group developed a significant response bias toward the now more frequently reinforced alternative. In contrast, the bias of the control group stayed near zero after an initial decline following the contingency change. The overall shift in bias was similar for the two groups. The behavior of both groups of children was sensitive to the asymmetric reward distribution and to the change in reward availability. Subtle group differences in response patterns emerged, possibly reflecting differences in the time frame of reward effects and sensitivity to reward exposure.

Broca's aphemia: an illustrated account of its clinico-anatomic validity.

OBJECTIVE: To present the case of a 54-year-old man with loss of speech, but with preservation of voluntary facio-lingual motility, language and other cognitive abilities (Broca's aphemia). METHOD: Observation of patient oral communicative abilities and general behavior, neuropsychological assessment and cranial computed tomography. RESULTS: Computed tomography showed a hyperdense lesion in the subcortex of the left precentral gyrus corresponding to Brodmann's area 6 and 44. Neuropsychological assessment confirmed that the major cognitive domains were intact. CONCLUSION: Our patient reiterates the validity of Broca's aphemia as a clinico-anatomic entity allowing us to portray it for the first time in pictures. From a neurobehavioral perspective, aphemia is related to apraxia rather than to aphasia, a fact that may have hampered the full grasp of its far-reaching implications for neurology and aphasiology.

Conversion technique from neuroendoscopy to microsurgery in ventricular tumors: Technical note.

BACKGROUND: Ventricular tumors represent a major neurosurgical challenge, making endoscopic approach an invaluable tool as it gained importance due to technological advances. Nevertheless, the method is not exempt of risk and limitations, sometimes requiring an open surgery. Thus, initial measurements must be adopted in order to simplify an eventual need for conversion to open craniotomy. METHODS: Here, we describe a series of 6 patients with ventricular tumors approached by neuroendoscopy where the conversion to microsurgery turned out to be necessary. Patients' average age was 59.5 years (39-75 years). Average tumoral size was 17.8 mm (15-21 mm). There were 2 cases of lateral ventricle subependymoma and 4 cases of third ventricle colloid cysts. A standard surgical incision was performed in the coronal direction, allowing lateral expansion to 10 cm. Moreover, the endoscopic burr hole was enlarged to a 5 cm craniotomy. A small enlargement of the endoscopic cortical access was performed to gain a transcortical microsurgical corridor to the ventricular cavity. The need for conversion arose due to high consistency of the tumor (3 cases), technical problems (2 cases), and cortical collapse (1 case). RESULTS: There was one case of cerebrospinal fluid fistula and infection and one case of transitory memory disturbance. In both the cases, we obtained a complete functional recovery. Clinical and radiological follow-up showed total tumor removal with no recurrences. CONCLUSIONS: The technique herein described was easy to perform, promptly bypassed the endoscopic limitations, and gathered excellent surgical results. The possibility of adapting the method to other tumor locations may be considered.

How Adaptive Learning Affects Evolution: Reviewing Theory on the Baldwin Effect.

We review models of the Baldwin effect, i.e., the hypothesis that adaptive learning (i.e., learning to improve fitness) accelerates genetic evolution of the phenotype. Numerous theoretical studies scrutinized the hypothesis that a non-evolving ability of adaptive learning accelerates evolution of genetically determined behavior. However, their results are conflicting in that some studies predict an accelerating effect of learning on evolution, whereas others show a decelerating effect. We begin by describing the arguments underlying the hypothesis on the Baldwin effect and identify the core argument: adaptive learning influences the rate of evolution because it changes relative fitness of phenotypes. Then we analyze the theoretical studies of the Baldwin effect with respect to their model of adaptive learning and discuss how their contrasting results can be explained from differences in (1) the ways in which the effect of adaptive learning on the phenotype is modeled, (2) the assumptions underlying the function used to quantify fitness and (3) the time scale at which the evolutionary rate is measured. We finish by reviewing the specific assumptions used by the theoretical studies of the Baldwin effect and discuss the evolutionary implications for cases where these assumptions do not hold.

A pilot study on effects of 4×1 high-definition tDCS on motor cortex excitability.

High-Definition transcranial Direct Current Stimulation (HD-tDCS) using specialized small electrodes has been proposed as a focal, non-invasive neuromodulatory technique. Here we provide the first evidence of a change in cortical excitability after HD-tDCS of the motor cortex, using TMS motor evoked potential (MEP) as the measure of excitability. Stimulation for 20 minutes at 1 mA with an anode centered over the hand area of the motor cortex and four surrounding return electrodes (anodal 4×1 montage) produced a significant increase in MEP amplitude and variability after stimulation, compared to sham stimulation. Stimulation was well tolerated by all subjects with adverse effects limited to transient sensation under the electrodes. A high-resolution computational model confirmed predictions of increased focality using the 4×1 HD tDCS montage compared to conventional tDCS. Simulations also indicated that variability in placement of the center electrode relative to the location of the target (central sulcus) could account for increasing variability. These results provide support for the careful use of this technique where focal tDCS is desired.

Community Genetics Reveal Elevated Levels of Sympatric Gene Flow among Morphologically Similar but Not among Morphologically Dissimilar Species of Lake Victoria Cichlid Fish.

We examined genetic structure among five species of Lake Victoria haplochromine cichlids in four island communities, using a full factorial sampling design that compared genetic differentiation between pairs of species and populations of varying morphological similarity and geographical proximity. We found that allopatric conspecific populations were on average significantly more strongly differentiated than sympatric heterospecific populations of morphologically similar species. Allopatric heterospecific populations of morphologically dissimilar species were most differentiated. Our work demonstrates that phenotypic divergence can be maintained and perhaps even evolve in sympatry despite considerable gene flow between species. Conversely, phenotypic resemblance among conspecific populations can be maintained despite geographical isolation. Additionally we show that anthropogenically increased hybridization does not affect all sympatric species evenly but predominantly affects morphologically similar and closely related species. This has important implications for the evolution of reproductive isolation between species These findings are also consistent with the hypothesis of speciation reversal due to weakening of divergent selection and reproductive isolation as a consequence of habitat homogenization and offers an evolutionary mechanistic explanation for the observation that species poor assemblages in turbid areas of the lake are characterized by just one or two species in each of a few morphologically distinct genera.

The self as a moral agent: linking the neural bases of social agency and moral sensitivity.

The human brain is inherently able to understand the world in moral ways, endowing most of us with an intuitive sense of fairness, concern for others, and observance of cultural norms. We have argued that this moral sensitivity ability depends on a sophisticated integration of cognitive, emotional, and motivational mechanisms, which are modulated by individual experience in different cultural milieus. Different lines of investigation on agency and morality have pointed to overlapping neural systems. Therefore, understanding the relationships between morality and agency may provide key insights into the mechanisms underlying human behavior in several clinical and societal settings. We used functional MRI to investigate the contribution of agency and of specific moral emotions to brain activation using action scripts. Results showed that emotionally neutral agency recruited neural networks previously associated with agency, intentionality and moral cognition, encompassing ventral and subgenual sectors of the medial prefrontal cortex (PFC), insula, anterior temporal cortex and superior temporal sulcus (STS). Compared to emotionally neutral agency, different categories of moral emotions led to distinct activation patterns: (1) prosocial emotions (guilt, embarrassment, compassion) activated the anterior medial PFC and STS, with (2) empathic emotions (guilt and compassion) additionally recruiting the mesolimbic pathway; (3) other-critical emotions (disgust and indignation) were associated with activation of the amygdala-parahippocampal and fusiform areas. These findings indicate that agency related to norm-abiding social behaviors of emotionally neutral scripts share neural substrates both with the "default mode" of brain function and with the moral sensitivity network. Additional activation in specific components of this network is elicited by different classes of moral emotions, in agreement with recent integrative models of moral cognition and emotion.

The moral affiliations of disgust: a functional MRI study.

Recent investigations in cognitive neuroscience have shown that ordinary human behavior is guided by emotions that are uniquely human in their experiential and interpersonal aspects. These "moral emotions" contribute importantly to human social behavior and derive from the neurobehavioral reorganization of the basic plan of emotions that pervade mammalian life. Disgust is one prototypic emotion with multiple domains that include viscerosomatic reaction patterns and subjective experiences linked to (a) the sensory properties of a class of natural stimuli, (b) a set of aversive experiences and (c) a unique mode of experiencing morality. In the current investigation, we tested the hypotheses that (a) the experience of disgust devoid of moral connotations ("pure disgust") can be subjectively and behaviorally differentiated from the experience of disgust disguised in the moral emotion of "indignation" and that (b) pure disgust and indignation may have partially overlapping neural substrates. Thirteen normal adult volunteers were investigated with functional magnetic resonance imaging as they read a series of statements depicting scenarios of pure disgust, indignation, and neutral emotion. After the scanning procedure, they assigned one basic and one moral emotion to each stimulus from an array of six basic and seven moral emotions. Results indicated that (a) emotional stimuli may evoke pure disgust with or without indignation, (b) these different aspects of the experience of disgust could be elicited by a set of written statements, and (c) pure disgust and indignation recruited both overlapping and distinct brain regions, mainly in the frontal and temporal lobes. This work underscores the importance of the prefrontal and orbitofrontal cortices in moral judgment and in the automatic attribution of morality to social events. Human disgust encompasses a variety of emotional experiences that are ingrained in frontal, temporal, and limbic networks.