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Mechanical sensitivity is commonly affected in chronic pain and other neurological disorders. To discover mechanisms of individual differences in punctate mechanosensation, we performed quantitative trait locus mapping of the response to von Frey monofilament stimulation in BXD recombinant inbred mice. Significant loci were detected on mouse chromosome (Chr) 5 and 15, indicating the location of underlying polymorphisms that cause heritable variation in von Frey response. Convergent evidence from public gene expression data implicate candidate genes within the loci: von Frey thresholds were strongly correlated with baseline expression of Cacna2d1, Ift27 and Csnk1e in multiple brain regions of BXD strains. Systemic gabapentin and PF-670462, which target the protein products of Cacna2d1 and Csnk1e, respectively, significantly increased von Frey thresholds in a genotype-dependent manner in progenitors and BXD strains. RT-PCR confirmed differential expression of Cacna2d1 and Csnk1e in multiple brain regions in progenitors and revealed differential expression of Cacna2d1 and Csnk1e in the dorsal root ganglia of the progenitors and BXD strains grouped by QTL genotype. Thus, linkage mapping, transcript covariance and pharmacological testing suggest that genetic variation affecting Cacna2d1 and Csnk1e may contribute to individual differences in von Frey filament response. The current study implicates Cacna2d1 and Ift27 in basal mechanosensation in line with their previously suspected role in mechanical hypersensitivity. Csnk1e is implicated for von Frey response for the first time. Further investigation is warranted to identify the specific polymorphisms involved and assess the relevance of these findings to clinical conditions of disturbed mechanosensation.
Two QTL, on chr 5 and 15, were identified for mechanosensation using the BXD RI strain panel. Candidate gene testing supports a role for Cacna2d1 (chr 5) and Csnk1e (chr 15) in mechanosensation variability
Duchenne muscular dystrophy (DMD) is a progressive muscle-wasting disorder, caused by mutations in the DMD gene and the resulting lack of dystrophin. The DMD gene has seven promoters, giving rise to multiple full-length and shorter isoforms. Besides the expression of dystrophin in muscles, the majority of dystrophin isoforms is expressed in brain and dystrophinopathy can lead to cognitive deficits, including intellectual impairments and deficits in executive function. In contrast to the muscle pathology, the impact of the lack of dystrophin on the brain is not very well studied. Here we study the behavioral consequences of a lack of full-length dystrophin isoforms in mdx mice, particularly with regard to domains of executive functions and anxiety. We observed a deficit in cognitive flexibility in mdx mice in the absence of motor dysfunction or general learning impairments using two independent behavioral tests. In addition, increased anxiety was observed, but its expression depended on the context. Overall, these results suggest that the absence of full-length dystrophin in mice has specific behavioral effects that compare well to deficits observed in DMD patients.
Cognitive inflexibility and anxiety in mdx mice in the absence of motor dysfunction or general learning impairments.
Loss-of-function mutations in progranulin (GRN) are a major autosomal dominant cause of frontotemporal dementia (FTD), a neurodegenerative disorder in which social behavior is disrupted. Progranulin-insufficient mice, both Grn+/− and Grn−/−, are used as models of FTD due to GRN mutations, with Grn+/− mice mimicking the progranulin haploinsufficiency of FTD patients with GRN mutations. Grn+/− mice have increased social dominance in the tube test at 6 months of age, though this phenotype has not been reported in Grn−/− mice. In this study, we investigated how the tube test phenotype of progranulin-insufficient mice changes with age, determined its robustness under several testing conditions, and explored associated cellular mechanisms. We observed biphasic social dominance abnormalities in Grn+/− mice: at 6–8 months, Grn+/− mice were more dominant than wild-type littermates, while after 9 months of age, Grn+/− mice were less dominant. In contrast, Grn−/− mice did not exhibit abnormal social dominance, suggesting that progranulin haploinsufficiency has distinct effects from complete progranulin deficiency. The biphasic tube test phenotype of Grn+/− mice was associated with abnormal cellular signaling and neuronal morphology in the amygdala and prefrontal cortex. At 6–9 months, Grn+/− mice exhibited increased mTORC2/Akt signaling in the amygdala and enhanced dendritic arbors in the basomedial amygdala, and at 9–16 months Grn+/− mice exhibited diminished basal dendritic arbors in the prelimbic cortex. These data demonstrate a progressive change in tube test dominance in Grn+/− mice and show that partial, but not complete progranulin deficiency disrupts this behavior in mice.
Biphasic tube test abnormalities in Grn+/− mice are associated with altered rpS6/mTORC2/Akt signaling and dendritic arbors in the amygdala and prefrontal cortex.
Recent genome wide association scans (GWAS) for reading and language abilities have pin-pointed promising new candidate loci. However, the potential contributions of these loci remain to be validated. In the present study, we tested 17 of the most significantly associated single nucleotide polymorphisms (SNPs) from these GWAS studies (p < 10−6 in the original studies) in a new independent population dataset from the Netherlands: known as FIOLA (Familial Influences On Literacy Abilities). This dataset comprised 483 children from 307 nuclear families, plus 505 adults (including parents of participating children), and provided adequate statistical power to detect the effects that were previously reported. The following measures of reading and language performance were collected: word reading fluency, nonword reading fluency, phonological awareness, and rapid automatized naming. Two SNPs (rs12636438, rs7187223) were associated with performance in multivariate and univariate testing, but these did not remain significant after correction for multiple testing. Another SNP (rs482700) was only nominally associated in the multivariate test. For the rest of the SNPs we did not find supportive evidence of association. The findings may reflect differences between our study and the previous investigations in respects such as the language of testing, the exact tests used, and the recruitment criteria. Alternatively, most of the prior reported associations may have been false positives. A larger scale GWAS meta-analysis than those previously performed will likely be required to obtain robust insights into the genomic architecture underlying reading and language.
FIOLA multivariate association results for candidate language and reading SNPs. WRF: word reading fluency, NWRF: nonword reading fluency, PA: phonological awareness, RAN: rapid automatized naming.
The developing fetus and neonate are highly sensitive to maternal environment. Besides the well-documented effects of maternal stress, nutrition and infections, maternal mutations, by altering the fetal, perinatal and/or early postnatal environment, can impact the behavior of genetically normal offspring. Mutation/premutation in the X linked FMR1 (encoding the translational regulator FMRP) in females, although primarily responsible for causing fragile X syndrome (FXS) in their children, may also elicit such maternal effects. We demonstrated in mice that a deficit in maternal FMRP results in hyperactivity in the genetically normal offspring. To test if maternal FMRP has a broader intergenerational effect, we measured social behavior, a core dimension of neurodevelopmental disorders, in males of FMRP-deficient dams. We found that male offspring of Fmr1+/- mothers, independent of their own Fmr1 genotype, exhibit increased approach and reduced avoidance toward conspecific strangers, reminiscent of "indiscriminate friendliness", or the lack of stranger anxiety, diagnosed in neglected children and in patients with Asperger's and Williams syndrome. Furthermore, social interaction failed to activate mesolimbic/amygdala regions, encoding social aversion, in these mice, providing a neurobiological basis for the behavioral abnormality. This work identifies a novel role for FMRP that extends its function beyond the well-established genetic function into intergenerational non-genetic inheritance/programming of social behavior and the corresponding neuronal circuit. As FXS premutation and some psychiatric conditions that can be associated with reduced FMRP expression are more prevalent in mothers than full FMR1 mutation, our findings potentially broaden the significance of FMRP-dependent programming of social behavior beyond the FXS population.
Maternal FMRP programs offspring sociability and alters activation of neural substrates of social interaction
Impulsivity is a multi-faceted construct that, while characterized by a set of correlated dimensions, is centered around a core definition that involves acting suddenly in an unplanned manner without consideration for the consequences of such behavior. Several psychiatric disorders include impulsivity as a criterion, and thus it has been suggested that it may link a number of different behavioral disorders, including substance abuse. Native Americans experience some of the highest rates of substance abuse of all US ethnic groups. The described analyses used data from a low coverage whole genome sequence scan to conduct a genome-wide association study of an impulsivity phenotype in an American Indian community sample (n = 658). Demographic and clinical information were obtained using a semi-structured interview. Impulsivity was assessed using a scale derived from the Maudsley personality inventory that combines both novelty-seeking and lack of planning items. The impulsivity score was tested for association with each variant adjusted for demographic variables, and corrected for ancestry and kinship, using EMMAX. Simulations were conducted to calculate empirical p-values. Genome-wide significant findings were observed for a variant 50 kb upstream from catenin cadherin-associated protein, alpha 2 (CTNNA2), a neuronal specific catenin, in the REG gene cluster. A meta-analysis of genome-wide association studies had previously identified common variants in CTNNA2 as being associated with excitement seeking. A second locus upstream of NEIL3 on chromosome 4 also achieved genome-wide significance. The association between sequence variants in these regions suggests their potential roles in the genetic regulation of this phenotype in this population.
We examined the relationship between 5-HTTLPR/rs25531 polymorphisms – related to serotonin-reuptake – and the ability to perceive and recognize faces and facial emotion expressions. We conducted structural equation modeling on data from 230 young adults, obtained by using a multivariate task battery. By additionally modeling fluid intelligence and immediate and delayed memory, we aimed to address the discriminant relationships of the 5-HTTLPR/rs25531 polymorphisms with socio-emotional abilities. We found a robust association between the 5-HTTLPR/rs25531 polymorphism and facial emotion perception. Carriers of two long (L) allelesoutperformed carriers of one or two S alleles. Weaker associations were present for face identity perception and memory for emotional facial expressions. There was no association between the 5-HTTLPR/rs25531 polymorphism and non-social abilities.
Identifying mechanisms through which individual differences in reward learning emerge offers an opportunity to understand both a fundamental form of adaptive responding as well as etiological pathways through which aberrant reward learning may contribute to maladaptive behaviors and psychopathology. One candidate mechanism through which individual differences in reward learning may emerge is variability in dopaminergic reinforcement signaling. A common functional polymorphism within the catechol-O-methyl transferase gene (COMT; rs4680, Val158Met) has been linked to reward learning where homozygosity for the Met allele (linked to heightened prefrontal dopamine function and decreased dopamine synthesis in the midbrain) has been associated with relatively increased reward learning. Here, we used a probabilistic reward learning task to asses response bias, a behavioral form of reward learning, across 3 separate samples that were combined for analyses (age: 21.80 ± 3.95; n = 392; 268 female; European-American: n = 208). We replicate prior reports that COMT rs4680 Met allele homozygosity is associated with increased reward learning in European-American participants (β = 0.20, t = 2.75, p < 0.01; ∆R2=0.04). Moreover, a meta-analysis of 4 studies, including the current one, confirmed the association between COMT rs4680 genotype and reward learning (95% CI −0.11 to −0.03; z = 3.2; p < 0.01). These results suggest that variability in dopamine signaling associated with COMT rs4680 influences individual differences in reward which may potentially contribute to psychopathology characterized by reward dysfunction.
COMT rs4680 Val allele carriers have blunted reward learning across 3 separate samples (European-American: n=208)
Face cognition, including face identity and facial expression processing, is a crucial component of socio-emotional abilities, characterizing humans as highest developed social beings. However, for these trait domains molecular genetic studies investigating gene-behavior associations based on well-founded phenotype definitions are still rare. We examined the relationship between 5-HTTLPR/rs25531 polymorphisms – related to serotonin-reuptake – and the ability to perceive and recognize faces and emotional expressions in human faces. For this aim we conducted structural equation modeling on data from 230 young adults, obtained by using a comprehensive, multivariate task battery with maximal effort tasks. By additionally modeling fluid intelligence and immediate and delayed memory factors, we aimed to address the discriminant relationships of the 5-HTTLPR/rs25531 polymorphisms with socio-emotional abilities. We found a robust association between the 5-HTTLPR/rs25531 polymorphism and facial emotion perception. Carriers of two long (L) alleles outperformed carriers of one or two S alleles. Weaker associations were present for face identity perception and memory for emotional facial expressions. There was no association between the 5-HTTLPR/rs25531 polymorphism and non-social abilities, demonstrating discriminant validity of the relationships. We discuss the implications and possible neural mechanisms underlying these novel findings.
We examined the relationship between 5-HTTLPR/rs25531 polymorphisms– related to serotonin-reuptake – and the ability to perceive and recognize faces and facial emotion expressions. We conducted structural equation modeling on data from 230 young adults, obtained by using a multivariate task battery. By additionally modeling fluid intelligence and immediate and delayed memory, we aimed to address the discriminant relationships of the 5-HTTLPR/rs25531 polymorphisms with socio-emotional abilities. We found a robust association between the 5-HTTLPR/rs25531 polymorphism and facial emotion perception. Carriers of two long (L) alleles outperformed carriers of one or two S alleles. Weaker associations were present for face identity perception and memory for emotional facial expressions. There was no association between the 5-HTTLPR/rs25531 polymorphism and non-social abilities.
Hedonic substitution, where wheel running reduces voluntary ethanol consumption has been observed in prior studies. Here we replicate and expand on previous work showing that mice decrease voluntary ethanol consumption and preference when given access to a running wheel. While earlier work has been limited mainly to behavioral studies, here we assess the underlying molecular mechanisms that may account for this interaction. From four groups of female C57BL/6J mice (control, access to two-bottle choice ethanol, access to a running wheel, and access to both two-bottle choice ethanol and a running wheel), mRNA-sequencing of the striatum identified differential gene expression. Many genes in ethanol preference quantitative trait loci were differentially expressed due to running. Furthermore, we conducted Weighted Gene Co-expression Network Analysis and identified gene networks corresponding to each effect behavioral group. Candidate genes for mediating the behavioral interaction between ethanol consumption and wheel running include multiple potassium channel genes, Oprm1, Prkcg, Stxbp1, Crhr1, Gabra3, Slc6a13, Stx1b, Pomc, Rassf5, and Camta2. After observing an overlap of many genes and functional groups previously identified in studies of initial sensitivity to ethanol, we hypothesized that wheel running may induce a change in sensitivity, thereby affecting ethanol consumption. A behavioral study examining Loss of Righting Reflex to ethanol following exercise trended toward supporting this hypothesis. These data provide a rich resource for future studies that may better characterize the observed transcriptional changes in gene networks in response to ethanol consumption and wheel running.
Striatal gene expression correlates with reduction in ethanol consumption due to wheel running.
Suicidal behavior and self-mutilation can be regarded as the expression of self-directed aggression and both are common in prison populations. We investigated the influence of externalizing behaviors, depressive symptoms, childhood trauma, 5-HTTLPR variants on self-directed aggression (N = 145) in a group of 702 male Italian prisoners. Participants were comprehensively evaluated, including for psychiatric disorders, impulsive traits, lifetime aggressive behavior [Brown-Goodwin Lifetime History of Aggression (BGHA)], hostility, violent behavior during incarceration, depressive symptomatology [Hamilton Depression Rating Scale (HDRS)], childhood trauma [Childhood Trauma Questionnaire (CTQ)]. Logistic regression analysis showed false discovery rate corrected independent main effects of externalizing behaviors: BGHA (P = 0.001), violent behavior in jail (P = 0.007), extraversion (P = 0.015); HDRS (P = 0.0004), Axis I disorders (P = 0.015), CTQ (P = 0.004) and 5-HTTLPR genotype (P = 0.02). Carriers of 5-HTTLPR high (LALA), intermediate (LALG, SLA) activity variants were more likely to have exhibited self-directed aggression relative to the low activity (LGLG, SLG, SS) variant: high/low: odds ratio (OR) = 2.3, 95% confidence interval (CI) 1.27–4.68, P = 0.007; intermediate/low: OR = 1.96, 95% CI 1.09–3.68, P = 0.025. The CTQ main effect was driven by physical abuse. There was no interactive effect of 5-HTTLPR and CTQ. Secondary logistic regression analyses in (1) all suicide attempters (N = 88) and (2) all self-mutilators (N = 104), compared with controls showed that in both groups, childhood trauma (P = 0.008–0.01), depression (P = 0.0004–0.001) were strong predictors. BGHA, violent behavior in jail predicted self-mutilation (P = 0.002) but not suicide attempts (P = 0.1). This study was able to distinguish differing influences on self-directed aggression between groups of closely related predictor variables within the externalizing behavioral domain. 5-HTTLPR had an independent, variant dosage effect.
Independent effects of 5-HTTLPR on self-directed aggressive behavior. Contingency analysis of self-directed aggressive behavior by 5-HTTLPR functional activity variants shows a variant dosage effect (L-R χ2 = 8.5, P = 0.014; Pearson χ2 = 8.1, P = 0.017). Triallelic 5-HTTLPR genotypes were grouped as low activity (SS, SLG, LGLG), intermediate activity (SLA, LALG) and high activity (LALA) variants.
Temperament is a key criterion in the selection of horses for both leisure and competitive riding to ensure optimal performance and safety. The Tennessee Walking Horse (TWH) is described as a calm, docile breed and is often used as a trail, show and pleasure horse. However, among horse owners and caretakers, there are anecdotes supporting familial and disciplinal typical behaviors and personalities. To investigate the contribution of genetics to temperament, we collected a behavior questionnaire, brief training history and identifying information for 276 TWH, as well as blood or hair samples for DNA. Factor analysis was conducted on the 20-item questionnaire for the set of 216 horses that met inclusion thresholds. Factor analysis identified four temperament factors in TWH: 'anxious', 'tractable', 'agonistic' and 'gregarious'. These four factors account for 64% of the total trait variance. DNA from 113 TWHs were selected and genotyped using the Equine SNP70 bead chip for three separate genome-wide association studies (GWAs) using the factor 1-anxious, factor 2-tractable and factor 3-agonistic scores as the phenotype. Quantitative association analysis identified significant candidate loci for each factor that warrant further investigation.
Genome-wide association analysis shows a significant peak on ECA13 for temperament factor 2-tractable in horses.
The mark/rouge test has been used to assess mirror self-recognition (MSR) in many species. Despite consistent evidence of MSR in great apes, genetic or non-genetic factors may account for the individual differences in behavioral responses that have been reported. We examined whether vasopressin receptor gene (AVPR1A) polymorphisms are associated with MSR-related behaviors in chimpanzees since vasopressin has been implicated in the development and evolution of complex social relations and cognition and chimpanzees are polymorphic for the presence of the RS3-containing DupB region. We compared a sample of DupB+/− and DupB−/− chimpanzees on a mark test to assess its role on social behavior toward a mirror. Chimpanzees were administered two, 10-min sessions where frequencies of mirror-guided self-directed behaviors, contingent actions and other social behaviors were recorded. Approximately one-third showed evidence of MSR and these individuals exhibited more mirror-guided self-exploratory behaviors and mouth contingent actions than chimpanzees not classified as passers. Moreover, DupB+/− males exhibited more scratching and agonistic behaviors than other male and female cohorts. Our findings support previous studies demonstrating individual differences in MSR abilities in chimpanzees and suggest that AVPR1A partly explains individual differences in MSR by influencing the behavioral reactions of chimpanzees in front of a mirror.
Male DupB+/− chimpanzees exhibited more scratching and agonistic behaviors than all other sex and genotype cohorts during a mirror self-recognition task.
Mutations in the X-linked cyclin-dependent kinase-like 5 gene (CDKL5) are associated to severe neurodevelopmental alterations including motor symptoms. In order to elucidate the neurobiological substrate of motor symptoms in CDKL5 syndrome, we investigated the motor function, GABA and glutamate pathways in the cerebellum of CDKL5 knockout female mice. Behavioural data indicate that CDKL5-KO mice displayed impaired motor coordination on the Rotarod test, and altered steps, as measured by the gait analysis using the CatWalk test. A higher reduction in spontaneous GABA efflux, than that in glutamate, was observed in CDKL5-KO mouse cerebellar synaptosomes, leading to a significant increase of spontaneous glutamate/GABA efflux ratio in these animals. On the contrary, there were no differences between groups in K+-evoked GABA and glutamate efflux. The anatomical analysis of cerebellar excitatory and inhibitory pathways showed a selective defect of the GABA-related marker GAD67 in the molecular layer in CDKL5-KO mice, while the glutamatergic marker VGLUT1 was unchanged in the same area. Fine cerebellar structural abnormalities such as a reduction of the inhibitory basket 'net' estimated volume and an increase of the pinceau estimated volume were also observed in CDKL5-KO mice. Finally, the BDNF mRNA expression level in the cerebellum, but not in the hippocampus, was reduced compared with WT animals. These data suggest that CDKL5 deletion during development more markedly impairs the establishment of a correct GABAergic cerebellar network than that of glutamatergic one, leading to the behavioural symptoms associated with CDKL5 mutation.
CDKL5 deletion impairs the establishment of a correct GABA- and glutamatergic cerebellar network, leading to locomotion abnormalities.
Human values and personality have been shown to share genetic variance in twin studies. However, there is a lack of evidence about the genetic components of this association. This study examined the interplay between genes, values and personality in the case of neuroticism, because polygenic scores were available for this personality trait. First, we replicated prior evidence of a positive association between the polygenic neuroticism score (PNS) and neuroticism. Second, we found that the PNS was significantly associated with the whole human value space in a sinusoidal waveform that was consistent with Schwartz's circular model of human values. These results suggest that it is useful to consider human values in the analyses of genetic contributions to personality traits. They also pave the way for an investigation of the biological mechanisms contributing to human value orientations.
This is the first demonstration of an association between human values and a specific genetic marker, the polygenic neuroticism score.
Genetic differences in acute behavioral responses to ethanol contribute to the susceptibility to alcohol use disorder and the reduction of anxiety is a commonly reported motive underlying ethanol consumption among alcoholics. Therefore, we studied the genetic variance in anxiolytic-like responses to ethanol across the BXD recombinant inbred (RI) mouse panel using the light–dark transition model of anxiety. Strain-mean genetic mapping and a mixed-model quantitative trait loci (QTL) analysis replicated several previously published QTL for locomotor activity and identified several novel anxiety-related loci. Significant loci included a chromosome 11 saline anxiety-like QTL (Salanq1) and a chromosome 12 locus (Etanq1) influencing the anxiolytic-like response to ethanol. Etanq1 was successfully validated by studies with BXD advanced intercross strains and fine-mapped to a region comprising less than 3.5 Mb. Through integration of genome-wide mRNA expression profiles of the mesocorticolimbic reward circuit (prefrontal cortex, nucleus accumbens and ventral midbrain) across the BXD RI panel, we identified high priority candidate genes within Etanq1, the strongest of which was Ninein (Nin), a Gsk3β-interacting protein that is highly expressed in the brain.
Behavioral genetics and genomic studies were identified as a novel locus modulating anxiolytic-like activity of ethanol in mice. We validated and fine mapped the most significant ethanol-anxiolytic QTL on Chr 12, termed Etanq1, and identified a highly promising candidate gene, Ninein. This is a very novel finding and because anxiety is considered a strong co-morbid factor for the development of alcohol use disorders and recidivism, we believe our studies are highly significant.
A growing body of evidence has shown that alcohol alters the activity of the innate immune system and that changes in innate immune system activity can influence alcohol-related behaviors. Here, we show that the Toll innate immune signaling pathway modulates the level of alcohol resistance in Drosophila. In humans, a low level of response to alcohol is correlated with increased risk of developing an alcohol use disorder. The Toll signaling pathway was originally discovered in, and has been extensively studied in Drosophila. The Toll pathway is a major regulator of innate immunity in Drosophila, and mammalian Toll-like receptor signaling has been implicated in alcohol responses. Here, we use Drosophila-specific genetic tools to test eight genes in the Toll signaling pathway for effects on the level of response to ethanol. We show that increasing the activity of the pathway increases ethanol resistance whereas decreasing the pathway activity reduces ethanol resistance. Furthermore, we show that gene products known to be outputs of innate immune signaling are rapidly induced following ethanol exposure. The interaction between the Toll signaling pathway and ethanol is rooted in the natural history of Drosophila melanogaster.
Reduction in Toll pathway activity decreases ethanol resistance, while activation of the Toll pathway increases resistance.
Phelan-McDermid syndrome (PMS) or 22q13.3 deletion syndrome is characterized by a variable degree of intellectual disability, impaired speech and language as well as social communicative skills and mild dysmorphic features. The SHANK3 gene is thought to be a major contributor to the phenotype. Apart from the syndrome-associated autistic features, symptoms from the bipolar spectrum can be discerned, in particular behavior instability and fluctuating mood culminating in a (hypo)manic state. In case of coincident major somatic events, a deteriorating course may occur. This study comprises seven adult patients (four females and three males; aged 21–44 years) with genetically proven PMS. Data from medical records were collected and extensive assessment of neuropsychological variables was performed to identify cognitive characteristics and their relation with psychopathology and treatment. All patients showed profound communication deficits and their developmental functioning ranged from 1.0 to 6.3 years. In addition, they had slow speed of information processing, impairment of attentional and executive functions and cognitive alexithymia. As to psychopathology, features from the affective and anxiety domains were prominent findings in these seven patients suggesting the presence of a bipolar spectrum disorder that could be effectively moderated with mood-stabilizing agents. Results are discussed in terms of the putative involvement of structural brain abnormalities, in particular cerebellar vermis hypoplasia and corpus callosum thinning and their cognitive and emotional sequelae. It is concluded that the treatment of 22q13.3-associated psychopathology should include prescription of mood-stabilizing agents in combination with individually tailored contextual neuropsychological measures.]]>
The dentate gyrus is one of the only two brain regions where adult neurogenesis occurs. Throughout life, cells of the neuronal stem cell niche undergo proliferation, differentiation and integration into the hippocampal neural circuitry. Ongoing adult neurogenesis is a prerequisite for the maintenance of adult hippocampal functionality. Bcl11b, a zinc finger transcription factor, is expressed by postmitotic granule cells in the developing as well as adult dentate gyrus. We previously showed a critical role of Bcl11b for hippocampal development. Whether Bcl11b is also required for adult hippocampal functions has not been investigated. Using a tetracycline-dependent inducible mouse model under the control of the forebrain-specific CaMKIIα promoter, we show here that the adult expression of Bcl11b is essential for survival, differentiation and functional integration of adult-born granule cell neurons. In addition, Bcl11b is required for survival of pre-existing mature neurons. Consequently, loss of Bcl11b expression selectively in the adult hippocampus results in impaired spatial working memory. Together, our data uncover for the first time a specific role of Bcl11b in adult hippocampal neurogenesis and function.
Loss of Bcl11b expression in the adult dentate gyrus impairs maturation of adult-born neurons and survival of pre-existing neurons.
Canine fucosidosis in English Springer spaniels is the only animal model of the neurovisceral lysosomal storage disease fucosidosis available for preclinical therapeutic trials. For this reason, it is crucial to identify critical time points in disease progression, and if there are particular lesions associated with specific aspects of neurologic dysfunction. Historical records of 53 canine fucosidosis cases from 1979 to 2009 containing a neurologic dysfunction score assessing motor, behavioral and sensory dysfunction were interrogated by statistical analysis. Motor and behavioral dysfunction scores assessing gait deficits and apprehensive behavior first significantly increased at 12–17 months, and increased at each 6-month interval thereafter. Sensory dysfunction scores, assessing hearing loss, balance and vision deterioration, did not significantly increase until 18–23 months, and coincided with a rapid decline in neurologic function. Regression analysis incorporating published neuropathology data, measured by image analysis, identified neuroinflammation and apoptotic cell death as significant informative predictors of increasing neurologic dysfunction. These findings indicate that the level of neuropathology required to induce consistent and conspicuous clinical signs in canine fucosidosis is reached by approximately 12 months of age in the absence of other disease processes. Significant association between neuroinflammation and apoptotic cell death also suggests that specifically targeting these lesions combined with enzyme replacement in future studies may reduce disease burden in fucosidosis. Overall, examining this historical clinical data to identify associations between the extent of neuropathology and degree of clinical dysfunction provides a useful reference tool for monitoring disease and evaluating therapeutic trials conducted in canine fucosidosis.
Neurologic dysfunction score in canine fucosidosis does not increase significantly until 12 months of age.
K+/Cl− cotransporters (KCCs) are known to be crucial in the control of neuronal electrochemical Cl− gradient. However, the role of these proteins in glial cells remains largely unexplored despite a number of studies showing expression of KCC proteins in glial cells of many species. Here, we show that the Caenorhabditis elegans K+/Cl− cotransporter KCC-3 is expressed in glial-like cells and regulates the thermosensory behavior through modifying temperature-evoked activity of a thermosensory neuron. Mutations in the kcc-3 gene were isolated from a genetic screen for mutants defective in thermotaxis. KCC-3 is expressed and functions in the amphid sheath glia that ensheathes the AFD neuron, a major thermosensory neuron known to be required for thermotaxis. A genetic analysis indicated that the regulation of the thermosensory behavior by KCC-3 is mediated through AFD, and we further show that KCC-3 in the amphid sheath glia regulates the dynamics of the AFD activity. Our results show a novel mechanism by which the glial KCC-3 protein non-cell autonomously modifies the stimulus-evoked activity of a sensory neuron and highlights the functional importance of glial KCC proteins in modulating the dynamics of a neural circuitry to control an animal behavior.
A potassium-chloride cotransporter functions in glia and modulates activity of a sensory neuron.


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Game Questions & Answers (Q&A) service lets you ask questions about video games for game consoles or PC games. So ask your Praetorians question for PC and get answers from real gamers or answer questions and share your insights and experience with the rest of the gaming community.
- Developer: Pyro Studios
- Publisher: Eidos Interactive
- Genre: Strategy Real-Time
- Release: Mar 10, 2003
- ESRB: Teen
Question & Answers
How to get past chapter 15
I am stuck on chapter 15 and need help to get past
Answer from: kurolhi
There are about 3 stucking chapter in praetorian. he who dares and when all hell break loose and the chapter before te last. in the chapter 15 , you need to start moving towards the blue coloured ally. one good way is save first and move the army and see where the enimies comes nd what type they are. then load and be redy for that. use the pickman for the first attack there with there bility of sitting with 2 row. behind them keep the archers with their most effective posture, for the next attack is from bow man standing high in the ground standing on the other side of Gabinnitio . kill them by the horse man eith the runnig speed. use the healer always after every attack. because the enemy army are large in number. after this attak must be similar way to the first time to those enemy marching outside from forest. meet the gabinnitio after that. after that build 2 defensive tower an put bowman. use a centurian. move him with care and go to any enimy area . they will follow him, but the advantage for him is that, no stamina is lost. so let them follow until u came across the defensive 2 towers. they will surely be beaten. remember not to keep any soldiers near the tower. or else they will attack them first. now repair the building nd do that again. clear the enimies around the blue ally. and meet them as soon as u can. or else they may be attacked. use this idea to therst of mission. have happy day
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Rob Lowe: Les Moonves Badgered Me Into Turning Down 'Grey's Anatomy'
Weeks after turning 50, Rob Lowe released his second memor "Love Life", detailing his stories as a husband and a father to two children as well as his career and romance. In the book, he reveals how he turned down what now becomes one of the most popular TV roles.
The "Killing Kennedy" star says he had verbally accepted the lead role on CBS' pilot "Dr. Vegas" when he got an offer to play Dr. Derek "McDreamy" Shepherd on "Grey's Anatomy". "After a week of negotiating, my deal was done, although not yet signed. It was then that I got an urgent call from the producers of a potential new show for ABC called 'Grey's Anatomy'," he recalls.
"I agreed to meet with the people making 'Grey's Anatomy'. I had read it and loved it-the writing was crisp, real and very entertaining-and it's always been a good idea to hear out talented people," he adds. "Grey's was a much better script; in fact, there was no comparison."
Though he loved the script, Lowe eventually turned down "Grey's" in favor of "Dr. Vegas". He admits that it's CBS boss Les Moonves who convinced him to do so. Moonves told the Golden Globe-nominated actor, "Let me tell you how it is. ABC is the lowest-rated network. ... At CBS we make hits."
Unfortunately, "Dr. Vegas" only lasted for five episodes. He, however, later became popular for his role as Robert McCallister on ABC's "Brothers & Sisters" before starring on NBC's "Parks and Recreation".
It wasn't the only juicy chance that Lowe missed. He reveals that Madonna once invited him backstage after a 1984 concert. They later hit a club together where she asked him to dance, but Lowe demurred and Madge left him and she never returned.
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- Is Nick Jonas Dating Lily Collins? Here's the Truth


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Have Pediatric Questions? Ask a Pediatrician.
He has constipation and belly pain. He had no fever or vomiting. He had an x-ray taken which showed the calcification. He was sent for an ultrasound and she could not find his appendix which she said is common in his age because it is so small and he had so much poo in his intestines. So they sent us home.
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The TSB Recorder
Do you have comments to share? Is there a story you'd like to see on the TSB Recorder?
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Latest post
The strength of working together
18 May 2016
Posted by Joe Hincke
Safety deficiencies must be uncovered, that's the imperative here at the TSB. Our investigators and teams of specialists are continuously working to do […]
Recent posts
Rail Safety – a collective effort
27 April 2016
Posted by Kirby Jang
The railway system has always fascinated me. Early in my career, while working in the private sector as a railway systems consultant, much of my work focused on the development of more effective processes for […]
The TSB then and now
7 April 2016
Posted by Johanne Ostiguy
When it was first created in 1990, the TSB's approach was one of "quiet professionalism". While the investigators and specialists conducted thorough investigations, leaving no stone unturned to uncover the root cause of […]
Canadian agency, global influence
10 March 2016
Posted by Brad Vardy
You often hear that TSB investigators are deploying to the scene of an occurrence in Canada, or that one of our investigation reports is being released. Our domestic activities are well known and publicised. But did you know […]
The right stuff
19 February 2016
Posted by Tony Gasbarro
Hi Tony, thanks for letting us interview you as part of the TSB's 25-year anniversary. Let me first start by asking, for how long have you been at the TSB? […]
Communication – at the heart of safety
8 February 2016
Posted by Marc-André Poisson
Over the years, the Marine Accident Investigators' Forum (MAIIF) and the International Maritime Pilots' Association (IMPA) have observed some shortcomings in communications between bridge team members […]
Older posts
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Print test
5 Written questions
5 Matching questions
- Dole
- the zimmerman teligram
- mussolini
- william howard taft
- cities
- a at the turn of the century,these grew faster than rural areas
- b the final major incident that promted the us to enter wwI
- c a hand out
- d theodore roosevelt's handpicked successor
- e the man ,known as IIpuca,who promised a second Roman Empire
5 Multiple choice questions
- Was the "red scare" a real threat?
- What attacks severaly crippled the US naval and air strangth?
- Who won the election of 1928?
- who helped Germany conquar Poland?
- the atomic bomb was dropped on wich cities?
5 True/False questions
Britain → the only allied power standing against Germany during the early part of 1941
Doughboys → Who amoung the Allies had their own units?
convoys → what did americans use in wwI to protect ships bound for europe?
ferdinand foch → Who develiped the esembly line?
Babe Ruth → Roosevelt's group of advisers


text:
Goalie Tyler Beskorowany was assigned from the Texas Stars to the Idaho Steelheads of the ECHL earlier this week so he could some more playing time. In his second game back with Idaho, Beskorowany stopped 39 of 40 shots last night in a 2-1 win over Utah. The Idaho Statesman's game story is here.
The Idaho Statesman also has a nice feature on Beskorowany's return to Boise, and you can read it here.
Nieuwendyk, other former Stars to play in alumni game
Stars GM Joe Nieuwendyk and several of his former teammates in Dallas are going to be lacing up the skates next month in an alumni game featuring players from the 1989 Stanley Cup Final between the Calgary Flames and Montreal Canadiens.
The game will be played on Saturday February 19 and it's part of the festivities surrounding the Heritage Classic game between the Canadiens and Flames, which will be played the following day.
Nieuwendyk was part of announcement about the alumni game yesterday in Calgary.
Also was on hand was former Star Brian Skrudland, a member of the 1989 Montreal team and a teammate of Nieuwendyk when the Stars won the Stanley Cup in 1999.
Here are the alumni game rosters as they stand now and you can pick out several players who played in Dallas over the years.
Calgary Flames: Colin Patterson, Dana Murzyn, Gary Roberts, Jamie Macoun, Jim Peplinski, Joe Nieuwendyk, Joel Otto, Lanny McDonald, Perry Berezan, Ric Nattress, Theo Fleury, Mike Vernon, Gary Suter (pending), Al McInnis (pending), Joe Mullen (pending).
Montreal Canadiens: Brent Gilchrist, Serge Boisvert, Benoit Brunet, Sergio Momesso, Mike Keane, Craig Ludwig, Kevin Haller, Guy Carbonneau, Bob Gainey, Russ Courtnall, Mike Lalor, Brian Skrudland, Ryan Walter, Kirk Muller, Rick Green.
The Flames official site has a nice feature on the game, including some quote from Nieuwendyk.


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Hokey Langan
Oct. 12, 2012
Every Friday, OhioStateBuckeyes.com features one of the many Ohio State social media outlets to follow. The Ohio State women's hockey team kicks off its home WCHA schedule this weekend, hosting nationally-ranked North Dakota at the OSU Ice Rink: OhioState_WHKY
The No. 10 Buckeyes are looking for the program's first 5-0-0 start this weekend, hosting No. 6 North Dakota. The squad is coming off an impressive sweep of then-No. 8/7 Minnesota Duluth on the road to open the WCHA season last weekend. It was Ohio State's first-ever sweep of the Bulldogs in the program's history.
Senior Paige Semenza leads the 4-0-0 Buckeyes with eight points on four goals and four assists. Junior Annie Svedin is second with six points (2g, 4a), while seniors Tina Hollowell, Minttu Tuominen and Hokey Langan each have five points.
Redshirt junior goalie Chelsea Knapp has a 1.25 goals-against average and was recently named the WCHA Defensive Player of the Week for her performances at UMD.
Friday and Saturday will each be promoting great causes at the OSU Ice Rink. On Friday, the team will be celebrating Sweetest Day and taking pop tabs and/or travel toiletries for Ronald McDonald House Charities. Saturday is Pink the Rink for Breast Cancer Awareness as pink buffs will be handed out with a post-game skate with some Ohio State players.
Other Ohio State women's hockey accounts to check out:
Twitter.com/NateHandrahan – Nate Handrahan, Ohio State women's hockey head coach
Twitter.com/BVondal – Ben Vondal, Ohio State Athletics Communications assistant
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Ohio State vs. Iowa B1G Tournament Game Postponed


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This is Michael. The most laid back cat ever! He likes any new lap and loves to pretend he never gets attention. His best friend was a tailless rat named Tess who he loved to play peek a boo with. Now he's usually cuddling his buddy simba.
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Italy will be without Mattia De Sciglio for their FIFA World Cup opener against England after the left-back suffered a spasm in his hamstring.
The AC Milan defender pulled up during training in Mangaratiba on Thursday after feeling a 'sting' in his left hamstring and his absence will force Cesare Prandelli into a backline reshuffle for Saturday's Group D clash.
The Italy coach could opt to move the experienced Giorgio Chiellini to full-back and pair Andrea Barzagli with either Leonardo Bonucci or Gabriel Paletta in the heart of defence.
Italy's chief doctor Enrico Castellacci confirmed De Sciglio will miss the game against England at a press conference.
"De Sciglio presented a left hamstring injury. Magnetic resonance imaging performed at Angra dos Reis showed swelling...De Sciglio will not be available for the opening match against England," Castellacci said.
But De Sciglio will remain in Italy's 23-man squad with Castellacci explaining the Milan full-back will be fit for his country's second group-stage match against Costa Rica on June 20.
"De Sciglio will stay with us. There are also no concerns for the second game against Costa Rica," the professor said.
Such late upheaval in Italy's back four could provide a cause for concern, with Barzagli having highlighted his unease over signs of Italian defensive frailties.
"In recent times, we have given away more goals than we wanted," the Juventus defender said.
"But the important thing is to give solidity to our defence.
"Against England, we will have to be very careful on dead-ball situations. They are very dangerous in the area and these are important details where we must not make mistakes."
0 - 1
35' C. Marchisio
(assist by M. Verratti)
D. Sturridge 37'
(assist by W. Rooney)
1 - 1
1 - 2
50' M. Balotelli
(assist by A. Candreva)
|J. Hart
|P. Jagielka
|G. Johnson
|L. Baines
|G. Cahill
|S. Gerrard
|J. Henderson
|R. Sterling
|W. Rooney
|D. Sturridge
|D. Welbeck
|Coach: R. Hodgson
|S. Sirigu
|A. Barzagli
|G. Paletta
|G. Chiellini
|M. Darmian
|D. De Rossi
|A. Pirlo
|C. Marchisio
|A. Candreva
|M. Verratti
|M. Balotelli
|Coach: C. Prandelli
for D. Welbeck 61'
for J. Henderson 73'
for D. Sturridge 80'
for M. Verratti 57'
for M. Balotelli 73'
for A. Candreva 79'
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*There is a double standard that comes with medical marijuana that causes a bit of confusion on the part of both the consumer and the general public.
No one would have considered the ramifications of medical marijuana use on those that have children. But now one father has to choose between his health and his child. (more…)
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Source: Total Wrestling Show
If the door is open for a WWE return: "I don't think that that door ever really closes as long as you haven't done something heinous. Yeah I would be open for a return in the future. Either place. I'm 29. I feel like I'm really coming into my prime as a Performers and I look forward to what lays ahead. Wherever it's going to be. It's going to be very fun, very exciting. The best is yet to come."
Training under Bill DeMott and the recent allegations against him: "He's old school. I was with Bill DeMott from the day Deep South opened until the day that it closed. I guess you could say that I'm a Bill DeMott guy. I went in there at 21 years old. Off the independents for a couple of years, very green and Jody had told Bill to make me quit. We would go in there and we would squat and bump. I would do 500-1000 squats a day & run 2 miles. Bill would make me do all of these things and then when they saw that I had heart and I had guts and that I deserved to be there, they really nurtured me and took me under their wing. I do think that sometimes in the past, in recent years, it's been too easy for guys. The sexual harassment thing, the allegations, seem like complete rubbish to me. I don't believe any of that. I think that those are people who are weak and they don't want to work hard. I'm more thankful for things that I have got and received from Wrestling than I would have been if it had just been handed to me coming in. I still have that old school perspective. If people disagree with that then so be it."
His career highlight: "I guess if it were one singular thing it would be when I main evented Smackdown against The Undertaker. Being 24 years and being able to pull off that match. I was really proud of that. There were so many things after that. Being a part of Wrestlemania. Working with Rey Mysterio in Mexico. Just a lot of different things. Going to Budokan Hall for the first time in Japan. Wrestling Sting on PPV. There's just a lot of stuff that were really cool things for me that I'll always remember. Like I said, I really think the best is yet to come. My prime years are ahead of me so there's going to be a lot more of that stuff."
His future: "I think the possibilities are endless. There are different things and places on the table. I am going to land on my feet. I always feel like that. When I came out of WWE I had a great year and a half working overseas and independently. Earning a great living, getting to see places I wouldn't have got to have seen in a big company. It'll be a big company sooner or later."
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America's Next Top Model 2003 | TV Show Watchlist
Season 8, Episode 2 The Girls Who Go to Prom
First Aired: March 7, 2007
The 12 remaining models get tips on how to walk a runway and participate in a prom-themed fashion show. Later, the women pose as high-school clichés in a photo shoot, before their number is reduced to 11 by the judges.
Next On TVSee all »
Thu May 26 8:01am
Dreckitude!OXYGNI'll Watch8 Watching
Cast & Details See all »
- Premiered: May 20, 2003
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- Premise: Aspiring models compete for a chance to break into the business in this runaway hit. Created and executive-produced by supermodel Tyra Banks, its formula of gorgeous women and men plus unappealing behavior make it enormously popular and one of the more int… (more)
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Deepening mystery around seized Vanuatu yacht
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South Pacific
Two men who crewed one of the world's most glamorous super-yachts, now detained in Vanuatu, have tried to flee to New Zealand, the Vanuatu Daily Post reports.
The 75-metre long Phocea was seized last week amidst speculation that it was running drugs, arms and was involved in money laundering.
Police also said at the time that two cabinet ministers were aboard but a tight news blackout has now been imposed.
The head of the country's transnational crime unit, Andrew Kalman, said in the wake of the initial raid on the yacht, he had been suspended from his job.
Last week Vanuatu police confirmed they launched the operation and arrested four people, including the skipper.
Police said the yacht had 16 crew members including a Samoan and a Tongan who had been taken into custody.
The two, named as Hiva Matiujaje and Michel Gangatadlakepa, have appeared in the Magistrate's Court on charges of obtaining Vanuatu citizenship by false and misleading statements. The prosecution attempted to have them remanded in custody, telling the court that both men had earlier been on bail but had been stopped trying to board a plane for New Zealand.
Neither name is a recognised one in Tonga or Samoa.
The men, pleading not guilty, will go to trial later this month.
The Daily Post said there was another raid on Phocea on Monday and items were confiscated.
It said police were also seeking the cabinet ministers for questioning.
Radio New Zealand International today reported that Kalman said his suspension is linked to the Phocea probe.
He said greed and special interests were behind his suspension, which he says has disappointed his immediate superior.
"I'm willing to find out who was the official complainant for my suspension, and I'm willing to take it further in relation to criminal defamation," he told RNZI.
Police refused to say anything further about the yacht, still moored in Vila Harbour.
The yacht had sailed from Italy through the Panama Canal, to Tonga and arrived in Vila nearly a fortnight ago.
Although Phocea is formally flagged to the land-locked duchy of Luxembourg, the Post says it is owned by a Thai national Anh Quan, who became a Vanuatu citizen this year. He is a principal in a company known as Amazonia Of The Pacific which is attempting to market agricultural produce internationally.
Phocea was built in France in 1976 and at the time was the largest yacht in the world.
In 1997, Lebanese businesswoman Mouna Ayoub purchased it for 5.5 million euros (NZ$8.4 million), part funding it by selling a 112 carat diamond. She is reputed to have spent US$20 million refitting the yacht.
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Winsome® Spectrum Pub Table with 2 Airlift Stools, Black
Item # WX2-196894SOLD OUT
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Winsome® Spectrum Pub Table with 2 Airlift Stools offer a stylish alternative dining spot.
Progressive design on this Spectrum Pub Table and pair of Airlift Chairs are perfect in your breakfast nook!
The 28" Table features a black MDF top that's built for years of use. Matching chairs are made with heavy-duty black ABS construction, and are padded and covered with black imitation leather.
Has stylish steel metal accents, too, for a look you'll love. Full 360° swivel and adjustable height with airlift mechanism makes the Stool even more comfortable!Details:
- Classic bucket-style seat
- Built-in metal footrest
- Easy assembly... all hardware included
- Chairs measure 19 x 18 2/3 x 43 1/2" and weigh 21 lbs. each. Table measures 28 3/4 x 28 3/4 x 40 1/8" and weighs 37 lbs.