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Metabotropic glutamate receptors (mGluRs) have well-established roles in cognition and social behavior in mammals. Whether or not these roles have been conserved throughout evolution from invertebrate species is less clear. Mammals have eight mGluRs whereas Drosophila has a single DmGluRA, which has both Gi and Gq coupled signaling activity. We have utilized Drosophila to examine the role of DmGluRA in social behavior and various phases of memory. We have found that flies that are homozygous or heterozygous for loss of function mutations of DmGluRA have impaired social behavior in male Drosophila. Futhermore, flies that are heterozygous for loss of function mutations of DmGluRA have impaired learning during training, immediate-recall memory, short-term memory, and long-term memory as young adults. This work demonstrates a role for mGluR activity in both social behavior and memory in Drosophila.
The Metabotropic glutamate receptors (mGluRs) in mammals have been shown to be involved in memory formation, long-term depression (LTD), long-term potentiation in mammals and linked to autism spectrum disorders (ASDs) in humans (Serajee et al., 2003; Mukherjee and Manahan-Vaughan, 2012). In mammals there are eight mGluRs, which are divided into three groups. Canonically, previous studies have established that group I mGluRs in mammals activate the Gq pathway, while group II and group III mGluRs activate the Gi/Go signaling pathway (Niswender and Conn, 2010; Chaki et al., 2012; Mukherjee and Manahan-Vaughan, 2012). However, there is accumulating evidence that in mammals, due to promiscuity of coupling to Gi and Gq, group II mGluRs may activate the Gq signaling pathway and induce LTD in a manner dependent on phospholipase C (PLC) and inositol trisphosphate receptor (InsP3R) activity (Huang et al., 1997, 1999a,b; Otani et al., 1999, 2002), and group I mGluRs are capable of activating Gi (Kreibich et al., 2004). In Drosophila there is only one mGluR, DmGluRA, which is coupled to Gi and Gq signaling (McBride et al., 2005; Pan and Broadie, 2007; Pan et al., 2008; Gatto and Broadie, 2009; Tessier and Broadie, 2012). Gi coupled signaling is able to engage and activate both the PI3K and ERK signaling cascades as well as increase glycogen synthase kinase-3 beta (GSK-3beta) activity and Gq mediated mGluR activation is also able to activate GSK-3beta activity (Fan et al., 2004; Huang et al., 2006; Beaulieu et al., 2009; Yuskaitis et al., 2010).
Drosophila DmGluRA expression has been demonstrated in the brain including expression in areas critically involved in short-term memory such as the antennal lobes (ALs) and mushroom bodies (MBs; McBride et al., 1999; Zars et al., 2000; Yu et al., 2004; Pesavento et al., 2008) and long-term memory in the MBs (McBride et al., 1999; Pascual and Preat, 2001) where expression is particularly heavy (Ramaekers et al., 2001; Pan and Broadie, 2007; Pan et al., 2008). More recently a detailed analysis of DmGluRA protein expression in the central complex has been published, a region of the brain where the expression of several other metabotropic receptors implicated in mammalian learning was found (Kahsai et al., 2012). The MBs in the insect are thought to be analogous to the mammalian hippocampus as first postulated from structural similarity to the human hippocampus in 1850 by the French physiologist and anatomist DuJarin (Dujardin, 1850; Davis, 1993, 2011; Busto et al., 2010; Skoulakis and Grammenoudi, 2006). Additionally, DmGluRA protein has been demonstrated to play a role in signaling at the presynapse of the NMJ in Drosophila and therefore could be similarly affecting signaling at the presynapse in the brain (Pan and Broadie, 2007; Pan et al., 2008; Banerjee et al., 2010).
The role of DmGluRA in cognition has been previously demonstrated in studies of Drosophila models of Fragile X syndrome and Alzheimer's disease. Fragile X is the leading inherited cause of intellectual disability and the leading known genetic cause of ASD. A fly model is based on loss of the of the fly dfmr1 gene, the ortholog of the human FMR1 gene, whose lack of expression leads to Fragile X syndrome. The Fragile X fly model has several behaviors in common with human symptoms including impairments in social interactions (Dockendorff et al., 2002) and cognitive impairments (McBride et al., 2005). Pharmacological blockade of the DmGluRA protein activity was able to rescue social interaction, immediate-recall memory and short-term memory in the Fragile X model representing the first time pharmacologic treatment rescued social impairments in an animal model of autism or memory impairments in an animal model of intellectual disability (Rubin, 1999b; McBride et al., 2005, 2012). Additionally, in this study, treatments initiated in development as well as those initiated in adulthood demonstrated efficacy in rescuing social interactions and memory.
DmGluRA has also been implicated in having a role in a Drosophila model of Alzheimer's disease that is based on mutations of the presenilin gene (McBride et al., 2010). The underlying nature of signaling alterations arising from the mutations in presenilin 1 or presenilin 2 genes that give rise to familial Alzheimer's disease (FAD) in humans are unclear (Saura et al., 2004; Walker et al., 2005; Qi-Takahara et al., 2005; Kumar-Singh et al., 2006; Sambamurti et al., 2006; De Strooper, 2007; Hardy, 2007; Isoo et al., 2007; Shen and Kelleher, 2007; Wolfe et al., 2007). Studies in model organisms indicate that the FAD-linked mutations lead to an impairment of presenilin 1 or presenilin 2 function (De Strooper, 2007; Kelleher and Purcell, 2008). This possibility suggests that some phenotypes associated with Alzheimer's disease, including age-onset cognitive loss, may be attributable to a reduction in overall presenilin protein activity levels. In the Drosophila Alzheimer's model young adult Drosophila (under 10 days of age, post-eclosion) have intact learning-during-training (LDT), immediate-recall memory and short-term memory, but have age dependent impairments in LDT and short-term memory at 30 days of age (McBride et al., 2010). Pharmacologic treatment with mGluR antagonists starting before cognitive impairments begin prevents cognitive impairment. Furthermore, treatment with mGluR antagonists starting after the onset of cognitive impairments reverses cognitive impairments in this model, indicating mGluR involvement in modulating synaptic plasticity well into adulthood (McBride et al., 2010). This indicates that in the Alzheimer's fly model, just as in the Fragile X fly model over active mGluR activity is contributing to memory impairment. More recently, under active mGluR activity has been implicated in phenotypes exhibited by tuberous sclerosis type 2 model mice (Auerbach et al., 2011). In spite of these findings, the involvement of DmGluRA in social interactions and memory in otherwise normal flies has remained unexplored in Drosophila. The purpose of this study was to examine the role of DmGluRA in social interactions and memory in Drosophila.
Social interaction can be examined in Drosophila in an ethologically relevant context by observing male courtship behavior directed toward female targets. Courting Drosophila males perform a characteristic sequence of behaviors: orienting toward and following the female, tapping her with his forelegs, vibrating one wing, licking her genitalia, and attempting to copulate (Bastock, 1955, 1956; Sturtevant, 1915). The percentage of time that the male spends performing any of these behaviors toward a target female during a defined period of time is referred to as the courtship index (CI; Siegel and Hall, 1979).
We first examined the ability of young adult (6–10 days post-eclosion) homozygous null DmGluRA112 flies to perform naïve courtship with virgin female targets as well as the DmGluRA2b, a precise excision control line. We found courtship behavior to be significantly impaired in the DmGluRA homozygous mutant flies, with the flies demonstrating almost no courtship activity (CIs of 3.2 ± 0.4), whereas the genetic background control flies demonstrated intact courtship behavior (CIs of 12.1 ± 0.8; Figure Figure1A1A). This demonstrated that the DmGluRA activity is required for social interaction since there was a significant impairment in naïve courtship behavior compared to the control strain.
The low courtship activity of the DmGluRA mutants prevented us from examining memory in the homozygous mutant mGluR flies. Therefore to explore a possible role of DmGluRA in memory, we examined courtship and memory in flies heterozygous for this mutation. To do this we crossed both the DmGluRA null mutant (DmGluRA112) and precise excision control (DmGluRA2b) lines to Oregon R flies. We found that in the heterozygous state there was still an impairment in social interaction of the DmGluRA112 heterozygous flies compared to the DmGluRA2b controls, 65.2 ± 5.3 vs 85.3 ± 6.2, again indicating a role for DmGluRA function in social interactions in Drosophila (Figure Figure1B1B). It should be noted that courtship in the OreR background is significantly higher than in the original background, mainly due to visual acuity differences in detecting motion, since the original background is white eyed. The OreR/OreR controls, the precise excision heterozygotes (DmGluRA2b/OreR) controls and the mutant heterozygotes (DmGluRA112/OreR) all had similar eye color. Furthermore, both control genotypes displayed similar levels of naïve courtship activity.
Although the heterozygous DmGluRA mutant flies displayed reduced naïve courtship activity, they still retained enough courtship activity to examine learning and various forms of memory using the conditioned courtship memory paradigm, an associative memory paradigm. In conditioned courtship, a male fly learns to modify his courtship behavior after experience with an unreceptive female (Siegel and Hall, 1979; Hall, 1994). Virgin females generally respond to a courting male by mating. However, recently mated females are unreceptive, display rejecting behaviors toward advances made by the male and have an overlapping but altered pheromonal profile that naïve males find less provocative than that of virgin female targets (Ejima et al., 2007). Normally, naïve male paired with a mated female target will initially court her, but his courtship activity soon decreases. This LDT is quantified, by comparing the CI during the first 10 min to the CI of the last 10 min period of a 1 h pairing with a previously mated female. In this paradigm wild-type flies typically show a ≥40% decrease in courtship activity (Joiner and Griffith, 1997; Kane et al., 1997). Hence, LDT is a form of behavioral plasticity but is distinct and separate from courtship suppression assayed after training, which is a form of associative memory (Tompkins et al., 1983; Ackerman and Siegel, 1986). When a male is paired with a virgin female after 1 h pairing experience with a mated female, his courtship remains depressed for 2–3 h (Siegel and Hall, 1979). This effect is not a general suppression of all courtship activity, because trained males do not modify their courtship of other pheromonally distinct targets (Ejima et al., 2005; Siwicki et al., 2005). After training with a mated female, memory is measured as a decrease in CI toward virgin females in trained males relative to naïve (sham trained) controls.
In Drosophila, five phases of memory have been elucidated by a combination of genetic and pharmacological dissection. There is an immediate-recall memory (immediate memory) at 0–2 min after training, short-term memory out to 1 h post-training, medium-term memory out to 6 h post-training, anesthesia-resistant memory out to 2 days post-training, and long-term memory lasting up to 9 days after training that appears to be dependent on protein synthesis (Skoulakis and Grammenoudi, 2006). Intact short-term memory is dependent on intact immediate recall. However, immediate recall and short-term memory are distinct from LDT. Therefore, intact memory can occur without LDT, and LDT can occur without post-training memory (Joiner and Griffith, 1997; Kane et al., 1997; McBride et al., 2005). Hence, in this study we chose to examine LDT, immediate-recall memory, short-term memory and long-term memory.
To assess LDT, a male fly was placed in a training chamber with a previously mated female for 1 h, and the amount of time the male spent courting in the initial 10 min interval was compared with the time spent engaged in courtship in the final 10 min interval. Heterozygous DmGluRA112 mutants display impaired LDT as young adults (Figure Figure22), similar to what he been previously observed in older DmGluRA112 mutant flies at 30 days of age (McBride et al., 2010). In contrast heterozygous DmGluRA2b controls and the OreR/OreR controls displayed intact LDT. This demonstrates a requirement for DmGluRA function in LDT.
To assess immediate-recall memory, a male fly was placed in a training chamber with a previously mated female for 1 h, and subsequently paired with a virgin female within 2 min of completing training. A lower CI compared to naïve trained (untrained) flies is indicative of memory. Heterozygous DmGluRA112 mutants display impaired immediate-recall memory as young adults, as they are not able to suppress their courtship upon subsequent pairing with a virgin female target (Figure Figure33). In contrast heterozygous DmGluRA2b controls and the OreR/OreR controls displayed intact immediate-recall memory. This demonstrates a requirement for DmGluRA function in immediate-recall memory.
To assess short-term memory, a male fly was placed in a training chamber with a previously mated female for 1 h, and subsequently paired with a virgin female 60 min after completing training. A lower CI compared to naïve-trained flies is indicative of memory. Heterozygous DmGluRA112 mutants do not demonstrate a suppression of their courtship upon subsequent pairing with a virgin female target, therefore they do not demonstrate short-term memory (Figure Figure44). In contrast heterozygous DmGluRA2b controls and the OreR/OreR controls displayed a suppression of courtship after training and therefore demonstrated short-term memory. This demonstrates a requirement for DmGluRA function in short-term memory.
Finally we examined if DmGluRA function was required for long-term memory (McBride et al., 1999; Banerjee et al., 2010). To assess long-term memory, a male fly was placed in a training chamber containing food with a previously mated female for 7 h, and subsequently paired with a virgin female 4 days after completing training (McBride et al., 1999; Banerjee et al., 2010). Again, a lower CI compared to sham trained (naïve-trained) flies is indicative of memory. Heterozygous DmGluRA112 mutants do not demonstrate a suppression of their courtship upon subsequent pairing with a virgin female target, therefore they do not demonstrate long-term memory (Figure Figure55). In contrast heterozygous DmGluRA2b controls and the OreR/OreR controls displayed a suppression of courtship after training and therefore demonstrated long-term memory. This demonstrates a requirement for DmGluRA function in the formation of long-term memory.
To ensure that the decreased courtship activity of the homozygous and heterozygous DmGluRA112 mutants was not the result of specific impairment in not being able to complete the various phases of courtship, we measured the percentage of flies that progressed through the stages of courtship. The homozygous DmGluRA112 mutants demonstrated the ability to progress through all of the stages of courtship in a 10 min testing period. Both the homozygous DmGluRA112 mutants and the homozygous control precise excision DmGluRA2b had a significantly lower percentage of flies progressing to the licking/attempted copulation stage compared to the flies that were crossed to the OreR background (p < 0.05 by chi square). However, the DmGluRA112 mutants and the homozygous control precise excision DmGluRA2b did not differ from each other in the percentage of flies that reached this final step (Figure Figure6A6A). Both of the heterozygous lines as well as the OreR background control reached similar percentages of achieving all stages of courtship (Figure Figure6A6A). Since both the homozygous and heterozygous DmGluRA112 mutant flies were observed to be capable of performing all of the steps of courtship, the lack of courtship activity does not appear to be secondary to some type of impairment that is rendering them incapable of completing all of the steps of courtship behavior. To ensure that the decreased courtship activity of the homozygous and heterozygous DmGluRA112 mutants was not the result of locomotor activity impairments, we examined locomotor function in the dishes utilized for the conditioned courtship testing (McBride et al., 2005, 2010). We did not find differences in spontaneous line crossing between homozygous or heterozygous DmGluRA112 mutant flies vs homozygous or heterozygous DmGluRA2b control flies or the OreR/OreR control flies (Figure Figure6B6B). Additionally neither the homozygous or heterozygous flies in any of the genotypes displayed gross impairments in olfaction or vision (Figures Figures6C6C,,DD).
Although a role for mGluRs signaling is well established in memory processes in mammals, the role of the only Drosophila mGluR, DmGluRA, has remained relatively unstudied with regard to social interaction or memory (Serajee et al., 2003; Mukherjee and Manahan-Vaughan, 2012). The expression of DmGluRA in the Drosophila brain is in areas critically involved in social behavior and memory including the ALs and the MBs, thus may contribute to these behavioral and cognitive processes (Ramaekers et al., 2001; Pan and Broadie, 2007; Pan et al., 2008).
Herein, we demonstrate that the DmGluRA activity is required for social interaction. We found that in the homozygous and heterozygous DmGluRA112 mutants have impairments in social behavior. This data fits well with the previous finding that a mutation in mGluR8 is linked to autism in humans (Serajee et al., 2003). Heterozygous DmGluRA112 mutants display impaired learning during training as young adults, thereby demonstrating a requirement for DmGluRA function in LDT, which may be analogous to working memory in mammals. Also heterozygous DmGluRA112 mutants display impairments in immediate-recall-memory, short-term memory, and long-term memory as young adults. These findings fit well with the known role of mGluRs in short-term and long-term memory formation in mammals (Mukherjee and Manahan-Vaughan, 2012). In our study the deficits in social behavior and memory did not appear to be caused by impairments in vision, olfaction, locomotion, or the capability to perform complicated coordinated motor tasks such as copulation or flight, thus DmGluRA function appears to be specific for social and cognitive tasks.
This study illustrates an evolutionarily conserved role of the mGluRs in synaptic plasticity and memory formation which is an important finding in the context of using lower organisms to model cognitive diseases such as Fragile X or Alzheimer's disease. Indeed, these are two disease models where antagonizing the Drosophila DmGluRA has been demonstrated to rescue social and/or memory impairments (McBride et al., 2005, 2010; Choi et al., 2010). It was in the Drosophila model of Fragile X that pharmacological blockage of DmGluRA protein function was first demonstrated to rescue social interaction, immediate-recall memory and short-term memory representing the first time pharmacologic treatment rescued social impairments in an animal model of autism or memory impairments in an animal model of intellectual disability (Rubin, 1999b; McBride et al., 2005, 2012). Additionally, it was demonstrated that treatments initiated in development as well as in adulthood demonstrated efficacy in rescuing social interactions and memory. The finding that adulthood treatments could ameliorate phenotypes associated with developmental disorders was paradigm shifting (Rubin, 1999a; State, 2010) and has now been demonstrated in other models of developmental disorders (Rubin, 1999c; Li et al., 2005; Guy et al., 2007). This strategy of decreasing mGluR activity to rescue cognition in the Fragile X model was later confirmed in mouse model of Fragile X by genetic and pharmacologic manipulation (Yan et al., 2005; Dolen et al., 2007; Choi et al., 2011) and has met with some early success in trials with Fragile X patients (Jacquemont et al., 2011). This demonstrates the important role of DmGluRA function not just in development, but also in adulthood, warranting additional studies.
In Drosophila or cell culture models of Alzheimer's disease antagonizing mGluRs has been efficacious in rescuing phenotypes associated with the models including cognitive impairment and Abeta secretion (Kim et al., 2010; McBride et al., 2010). Furthermore, both agonist and antagonists of mGluRs are under development for the treatment of depression in humans (Chaki et al., 2012; Mukherjee and Manahan-Vaughan, 2012). The current work along with the extensive mammalian literature on the involvement of mGluRs in memory illustrate that caution should be observed when looking at the mGluRs as receptors to modulate for the rescue of disease specific symptoms, because they may have unwanted effects on other aspects of cognition.
At first pass our results demonstrating that reduction of DmGluRA activity negatively impacts social behavior and cognition may seem counterintuitive, because antagonism of this receptors signaling can enhance memory in specific disease models. First, we have previously found that treatment with mGluR antagonists does not enhance memory in wild-type flies, indeed they impair memory and social activity (McBride et al., 2005; Choi et al., 2010). Also, toward this point it is important to keep in mind the way the molecular signaling occurs during memory formation. DmGluRA is predominantly coupled to Gi, thereby suppressing cAMP signaling. There is well documented literature in the fly field that indicates that cognition is impaired if cAMP levels are either too high, or too low. The dnc mutant has severe memory impairments and results from too much cAMP. The dnc mutation would be analogous to the DmGluRA mutants, with too much cAMP. In contrast the rut mutation leads to too little cAMP and also results in memory impairment (Skoulakis and Grammenoudi, 2006). This would be analogous to the fly models of Alzheimer's disease and fragile X syndrome, where the problem is too little cAMP and it is corrected by treatment with mGluR antagonists which should correct the cAMP to a level were normal memory can occur (McBride et al., 2005, 2010; Choi et al., 2010, 2011).
In conclusion, this work demonstrates that in Drosophila, just as in mammals, proper DmGluRA function is required for social behavior and various aspects of cognition including LDT, immediate-recall memory, short-term memory, and long-term memory.
Virgin male flies were collected under ether anesthesia within 4 h of eclosion. Virgin XX, y, f (attached X) females were collected on the day of eclosion and kept in food vials in groups of 10–15. Flies were aged in a 12 h light/dark cycle before behavioral training and testing. All testing was performed during the relative light phase. Mated females were 5 days old and observed to mate with a male the night before training. The virgin females that were used as targets were 4 days old (Joiner and Griffith, 1997; McBride et al., 1999, 2005).
For courtship behavior testing, males of the appropriate genotypes were collected within 4 h of eclosion and kept in isolation before testing. All flies were kept in 12 h light/dark cycles at 25°C and 50–75% relative humidity and were aged 6–10 days post-eclosion before training. All male subjects were transferred to fresh control food the day before testing (McBride et al., 1999, 2005, 2010). Male flies were assigned to random groups for behavior training and testing, which was performed blind (Siegel and Hall, 1979; Kane et al., 1997; McBride et al., 1999). The total amount of time a male was engaged in courtship activity while paired with an unanesthetized target female during a test period of 10 min or until successful copulation occurred was scored. A CI was calculated as the percentage of total observation time spent courting (Siegel and Hall, 1979). Testing of naïve courtship, LDT, immediate-recall and short-term memory were performed as previously described (McBride et al., 1999, 2005). For the naïve courtship analysis, the male was sham trained for 1 h in the training chamber without the addition of the female. The male was then transferred to the mating chamber containing a virgin female. Males were monitored for courtship activity that included orienting, following of the female, wing extension and vibration, tapping of female with his foreleg, genital licking and attempted copulation for a period of 10 min, or until copulation occurred.
Measurement of immediate-recall was made by pairing a naïve male with a non-receptive female for a single 1 h training session and then placing him in a second chamber with a receptive female within 2 min of completing training. Short-term memory was assessed by taking a male that had been trained with a non-receptive female for 1 h and placing him in isolation for 1 h before pairing with a virgin, receptive, female. At least 16 animals were tested for each genotype during analyses of naïve courtship, learning during training, immediate recall, short-term memory, and long-term memory.
The training paradigm for assessment of long-term memory is derived from McBride et al. (McBride et al., 1999; Banerjee et al., 2010). Males were paired with a non-receptive female for seven continuous hours and then kept in isolation for 4 days before testing. Sham-trained males were treated identically, except for the exposure to the training female. The observers were blind to the genotypes of the animals for all courtship studies (Banerjee et al., 2010; Sidyelyeva et al., 2010). Locomotor, olfaction, and visual acuity testing was done as in the study by McBride et al. (Griffith et al., 1993; McBride et al., 2005; Orgad et al., 2000).
The Drosophila strains were cultured as in the study by McBride et al. (2005). The DmGluRA mutant and control lines used during this study are white eyed and have been previously described and utilized, they are the previously described null allele of DmGluRA (DmGluRA112) and precise excision wild-type allele (DmGluRA2b) that provides an appropriate background control for the null allele (Bogdanik et al., 2004). Heterozygous versions of the mutant and control were obtained by crossing males to Oregon R virgin females. Heterozygous DmGluRA112 and DmGluRA2b F1 males were selected from the resultant progeny (McBride et al., 2010). The DmGluRA locus is on the 4th chromosome.
Courtship index of tested males were subjected to arcsin square root transformations to approximate normal distributions since not all of the sets of data were normal distributions, as is common in conditioned courtship data sets (Joiner and Griffith, 1997; McBride et al., 1999, 2005). For statistical comparison between the genotypes and treatments, Two Way ANOVA was used for genotype and treatment, with genotype resulting in a p value of 0.0001 and treatment resulting in a p value of 0.0001. The interaction p value was 0.0001. The post hoc analysis used for the comparison for the figures was the Bonferroni analysis (Villella and Hall, 1996; McBride et al., 1999, 2005; Ejima et al., 2007). For the figures we have placed asterisks according to the post hoc Bonferroni analysis since this is demonstrating memory or no memory within a specific genotype according to the provided experience (treatment) of the flies (Villella and Hall, 1996; McBride et al., 1999, 2005, 2010; Ejima et al., 2007). For line crossing experiments standard student t test was used and for Figure Figure66 binning analysis, olfaction and vision studies chi squared analysis was performed (McBride et al., 2005). All statistics were performed using both Statview 3.0 and Prism 5.0.
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
This work was supported by the FRAXA Research Foundation with grants to (Sean M. J. McBride, Catherine H. Choi, Thomas A. Jongens), National Institutes of Health (NIH) Grants GM086902 (Thomas A. Jongens), an Alzheimer's Disease Center pilot award from NIH Grant AG-10124 (Thomas A. Jongens), as well as an Alzheimer's Disease Research pilot award, a program of the American Health Assistance Foundation (Thomas A. Jongens). Thomas V. McDonald and Sean M. J. McBride were supported by a grant from Autism Speaks, and Eric Koenigsberg was supported by a summer research fellowship grant from The National Fragile X Foundation. The Albert Einstein College of Medicine MSTP grant funded Sean M. J. McBride. Sean M. J. McBride was also supported by an R25 MH060490 (Clinical Research Scholars Program in Psychiatry). We thank Evan Braunstein, Joseph Hinchey, Sean Campbell, Myles Akabas, Nancy Carasco, Peter Davies, and Kami Kim for critical comments during this project. We thank Oliver Schipper for help with fly husbandry. We thank Susan Horwitz and Charles Rubin for critical comments and generously providing space and resources.
PubMed Central Canada is a service of the Canadian Institutes of Health Research (CIHR) working in partnership with the National Research Council's national science library in cooperation with the National Center for Biotechnology Information at the U.S. National Library of Medicine(NCBI/NLM). It includes content provided to the PubMed Central International archive by participating publishers.


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Ukraine is burning. Ongoing fighting between government forces and insurgents in eastern Ukraine have left more than 2,000 Ukrainian civilians dead and some 300,000 displaced, according to AP. Meanwhile, in large swathes of the Donetsk and Luhansk regions, those who remain lack food, running water, and electricity. If the West is serious about preventing things deteriorating further it needs to find a new approach – and one that makes Russia part of the solution.
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crow's feetnoun [ plural ]uk /ˈkrəʊz fiːt/ us /ˈkroʊz fiːt/
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A platonic relationship or emotion is loving but not sexual.


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Perfume Bottles in Ancient Occasions The enticing scent in the best perfume as well as the eye-catching elegance in the bottle has become an important part of globe culture considering that ancient occasions. The oldest identified perfume bottles came in the Ancient Egyptians. They crafted containers manufactured from stone and alabaster which kept the perfume interesting, and prevented the liquid from leaking. The Ancient Egyptians also utilised clay pots they sculpted into styles of individuals and animals. Perfume was an essential a part of the Egyptians planning of your dead. A lot of of those ancient perfume bottles are actually located in tombs. Containers are actually located made from wood with diverse compartments for each perfume. Glass bottles were also a a part of ancient perfume history. They date every one of the way back on the fifteenth century BC. 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Go for any relatives game of mini golf, take a journey on the seashore, get a picnic to the park, or go for an ice cream or coffee just the two of you. Taking a walk together may be very simple, but it exhibits your Dad which you worth time with him and therefore are generating it a priority. Take the chance to ask your Dad concerns about what it had been like for him expanding up, what important changes he is noticed in his lifetime, and what guidance he can supply you with. five. Present Vouchers Everybody says that vouchers and present cards are boring and it feels such as the giver hasn't offered time or believed in to the present. Alter the typical perception! Give tickets to his favourite sporting occasion, but give a small piece of his team's merchandise with it so he has a thing to unwrap over the day. Make him go on the treasure hunt for his gift card. Or give him a present card and state that you'd like to feature him when he spends it and just hang out with him and have a coffee collectively when you are both completed. The top rated keys to offering great Fathers Day presents is really in producing some thing distinctive and tailored exclusively for the Dad's interests. Believe outdoors the box, and also have enjoyable producing recollections this Fathers Day!So why must you take into account looking at Honest Trade presents this Christmas time? Actually precisely what is Honest Trade? The Cartier Firm is usually a subsidiary of Compagnie Financi??re Richemont SA. Earlier than 1964, the corporate was owned and managed by the Cartier loved ones of French jewelers. The Cartier family was renowned as artists of elaborate jewelry, commissioned by royalty from all corners of your earth. The brilliance of their system and exotic substitute of jewel ensembles designed such grand goods since the "Bestiary" (illustrated by the Panth??re brooch in 1940, designed for Wallis Simpson,) the diamond necklace that Yadavindra Singh Maharaja of Patiala wore in 1904, and naturally the well-known Santos wrist regarder, produced for your Brazillian aviator Alberto Santos Dumont. The Cartier La Dona montres possess a single flaw. They could be mainly highly-priced. Acquiring a replique Cartier La Dona timepiece is really a big quantity of dollars substantially less. Even though a replique, there exists however a distinguishable degree of large high quality from other timepieces. The clarity of your glass glimmers flawlessly and like a consequence from the dimension on the dial it is in fact effortlessly Replique Montre examine in virtually any lights circumstance .cartier mens love bracelet
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From the occasion that you are a follower of acquiring quite a few the jewelry following which creating the real cartier jewelry, you could possibly also get the jewellery which is finished. That you are ready to straight pick out by indicates from the extremely several the cartier jewellery readily accessible as an example bracelets, necklaces and so forth., you can invest in these jewelry and send for your household members. The real organization releases fresh kinds associated with charms each and every solitary yr plus some while in the aged necklaces are removed by signifies through the manufacture assortment. You will discover numerous new necklaces that are at this time made readily available to individuals in recent times inside your prior couple of associated with quite a few as for which necklaces that are at the moment stopped, they've turned into memorabilia which might be slowly expanding within their definitely well worth evening greatest following day time. inside situation you should collect these stopped necklaces, it's the bulk of substantial that you just just acquire all of them the actual second more than time these are now removed through means in the production assortment prior to to they are going up in require and value.The hat was a very important detail of each woman of 'model of S during long centuries. In fact, a couldn of girl 't is called elegant if it didn 'use of T a hat just fifty years ago. Which was you the last time where saw an astonishing hat of wearing of elegant woman (except the political meetings)? In way or other nowadays of the hats are forgotten. The only hats which you can note in streets are hats of the sun. The various colors are with the mode each season, but should rather seek that to you (or several) these complements your skin. If the color of hat makes your color of face to seem strange, him 's better to still choose. You are going to possess a decision of a number of semi-precious stones and lots of pick out their birthstone for your leading of your ring. There may be an engraving additional beneath the stone, which adds a great deal of depth towards the ring layout. Think about the precious metal that can be applied for the ring when selecting your stone, also. You'll find a few of the stones that search superior with gold than silver and vice versa. Make sure you buy the correct dimension. While you might imagine that this goes without the need of saying, it ideal to use whatever ring sizer is supplied within the website youe ordering on. You just print and minimize the sizer out and youl know that youe ordering the proper size. Now, youl move onto probably the most significant element of ring design and style ?the personalization. You will have your selection of in excess of one hundred emblems, symbols, and insignias. Look at working with the crest to the branch of service around the prime with the ring, both below or with no precious stone. A unit or division insignia is usually a fantastic choice for one side with the ring. Another could possibly attribute an aircraft or ship the military member is assigned to. Possibly a specific date or initials will be a terrific choice or an insignia for a particular military campaign.A handful of examples are going to be described right here to display what a great choice they have. The 1st view could be the T Touch Professional titanium. That is for guys, and it can be made from a powerful titanium. This view is additionally waterproof helping to help keep it practical for really quite a few many years. The dial is usually a black carbon shade which offers it a a lot more stylish physical appearance. The 2nd observe they have would be the Seatouch watch. These are pleasurable pieces which appear good on these informal days. They've got precisely the same high-quality, however the band is manufactured from leather. It has a vibrant color which seems to be terrific on individuals summertime days. This observe is waterproof as well. The third example for males will be the through the Silent collection. These watches work effectively using the qualified seem since the band to the observe is leather. The face in the watch is encased in the stainless steel shell. The dial is really a black carbon which offers it additional fashion. Switching to collections made specially for ladies, the Equit collection is often a exceptional alternative for just about any elegant event. This watch includes a encounter that is an extended rectangular form. It is actually manufactured from stainless steel. And also the principal cause of giving the rings for very affordable costs is they've to shell out reduced overheads and operation value. Another area from exactly where you are able to invest in your engagement ring for very affordable charges is pawn outlets or thrift shops. You'll be able to also buy 2nd hand engagement rings and might simply save money on your ring. By surfing the web you are able to find auction sites giving 2nd hand engagement rings for price reduction costs. But in situation for those who don't wish to purchase your ring on the internet then you definitely can seem for wholesale community jewelry outlets. In these stores you can uncover the identical rings for wholesale prices which it is possible to find in regional jewelry shops for high priced charges. Hence for those who certainly are a sensible buyer then it can be clear that you will buy your ring from the wholesale jewellery shop because there is absolutely no ought to devote additional when you can get the same piece at fewer costs. One more most effective method to get affordable engagement rings should be to select much less expensive metals to your ring. Often bear in mind if you want to save money on your ring then you definately need to steer clear of platinum or yellow gold. The SIZES are by far the easiest things to look for. Pretty much in just about every replica on the earth, you can discover that whether or not the engravings/markings are appropriate, the sizes are normally off slightly. Either the Roman numeral markers are smaller sized, the luminescent markers are smaller sized, or typically, the minute/hour hands aren't precisely the same size and shape. One big problem identified on most replicas is every single replica may be somewhat diverse. This is regardless of regardless of whether it can be a Swiss, Japanese, or even Asians. Some are produced with much more markings than other folks; some are produced with a lot more gold plating than other people Fake Rolexes and fake Panerai are incredibly well-known currently. I advise all my readers at www.replicawatchesplus.com to perform their investigate ahead of obtaining. A lot of the fake Rolexes are very shut, and need mindful review to identify. Should the cost is unrealistically very low, consumers beware! Replica Watches demonstrates you the ins and outs of counterfeit watches. Cartier can be a title synonymous with elegance and good quality. The story of how Cartier watches came to get is definitely an exciting 1. Today, Cartier is a title that characterizes class and sophistication. For that reason, you will discover models that go for around $67,000. Not all watches are this pricey while most Cartier can have a hefty cost tag. They didn start out very in this way, even so. Cartier watches began in 1847 when Louis-Francois Cartier took ownership of his master jewelry workshop at 29 Rue Montogueil in Paris. Later on, in 1899 this deal with would transform to 13 Rue de la Paix. This is often the street known for grandeur and sumptuousness in Paris. These exquisite watches would expand a rich clientele and fan base around the globe. During the early years of 1900, an aviator good friend on the Cartier household complained with regards to the techniques of reading the time from a fob view although flying. Cartier produced an answer inside a proto-type wristwatch. This was a common trouble of aviators with the time. Cartier was promulgated from the Prince of Wales in 1902 because the Jeweler of Kings for King of Jewelers. Cartier used metals this kind of as platinum, steel, titanium and colorful gold (yellow, white, pink) to make their watches and they even now do that nowadays. Cartier creates watches of each size and design in the mini to your further large, the round to square and much more.
The blending of these two styles from two different eras has gone down effectively using the youngsters of right now and so they go crazy for this kind of stuffs. Considering that it truly is feasible to reproduce any dressing design irrespective with the period in the course of which the identical was worn the medieval variations have been perfectly accommodated inside of the modern day types. The end result is we have got wonderful dresses that look really appealing and individuals create a beeline to buy those. At present people today of all age groups obtain medieval dresses. Anything which has been unique throughout the medieval instances is reproduced right now and such clothing have numerous takers. When men and women get started wearing those costumes they turn into extremely popular and even more people today come to obtain such costumes. So when designers re-create the fashion in the yesteryear's they're sure that individuals will definitely value these. In a different sense men and women of this modern day age wear such clothing to express their attachment on the tradition and design of their ancestors. The medieval era lasted from 1200 to 1550. Each men and gals of that period wore long tunic fashion garments manufactured from wool because it was by far the most popular fabric. Any outlet that is certainly totally licensed to accomplish this may often be marketing authentic Glasses and consequently you will usually know exactly what you happen to be receiving. Certainly, it truly is attainable for you personally to get some far more very affordable and nevertheless real Plain Glasses in the event you shop all around somewhat bit, especially on the net.
The watches have lots of possesses in various value ranges. If you're curious in dealing wholesale watches then you can make use of this as a guide. You may purchase them from a wholesale watch supplier so you can get massive discount rates. You may also try a drop shipping corporation so you wouldn't must difficulty about stocking your orders and dispatching them for your consumers. With drop shipping, a retailer can protect income because you do not really need to acquire the item until eventually it truly is traded. You'll find a great deal of of appropriate and responsible drop shipping companies from Salehoo's directory list. Basically, there are actually alot of positive aspects you could get from contracting up as being a Salehoo member. For one particular, you may get a full admission to their checklist of ideal suppliers. You can even come across the fundamentals of drop shipping. You may also get incentive books about how to obtain the soundest wares to promote on the web and a lot more.Fathers Day is often a major day for each Dads and youngsters alike. Year right after year however, it may get a bit tedious to figure out what precisely to do to produce this Fathers Day an extra unique day. Here are some top tricks to make your Dad's day an additional memorable and exclusive day: 1. Get it done Homemade What Dad does not like sweet factors? Make him his favourite dessert or snack. If he loves to barbeque, why not invest in a blank apron, dig out the material paints, and design a bbq apron your Dad are going to be proud of? As an alternative to purchasing a card to go together with your existing, why not make your own? Make it unique with pop-ups, fancy models, and maybe even a pocket of vouchers of points you might do for Dad this Fathers Day!cartier love bangle dupe
Using the coming Breitling Replique Montre of 2012, Rado Montres have continued to enhance on their Montre style. Rado is known for his or her highly distinctive and fabulous designs. In an effort to continue this best reputation in their own taste of Breitling Navitimer Replique design, they have not disillusioned the clients with their new styles therefore persevering with their track record for their leading notch style. Using the help of advanced technology and also the utilization of expensive resources, Cartier Montres are nicely received by people for large good quality, leading craftsmanship and art beauty. Specifically Cartier jewelry Montres, they are all luxury with gems and diamond charm. Using the top quality craftsmanship and production, great pricing and excellent client provider why would you would like to obtain from anywhere else? You might get fast delivery on any item which you purchase as properly because the assure which you or your beloved one particular will love any with the IWC Portuguese Montres that you simply choose on. the Vacheron Constantin Tourbillon may be an uncommon and very detailed design with a fantastic style that makes it synonym of ease and comfort but also elegance. For jewellery purposes it is often alloyed with not less than one particular extra metal to make sure power and durability. 24 karats gold is the purest form of gold but the majority of the rings produced from gold are both 14 karats gold or 18 karats gold. If you will see a ring is marked as 14 karats gold you then realize that it is actually roughly 58.3 percent gold, whereas an 18 karat ring is 75 percent pure gold. An 18 karats gold ring is extra susceptible as there is more gold in it and as a result it is weaker rather than durable evaluating to 14 karats gold. A lot more gold you select for the ring could be the weaker ring you select. In the option of gold you will find also quite a few variations as white gold and yellow gold. If you need a ring that will have a white touch or far better to say a silver lining then white gold is definitely the solution. It's going to include a silver tone to your ring. White gold is simply yellow gold which has been handled to seem white (or silver). Now in an effort to make a yellow gold into white gold aspects like nickel, palladium, or silver are added. Nickel can give yellow gold a good white gold physical appearance. However the challenge with this is it is actually allergic and consequently can cause skin troubles. So it will not be harmless to dress in a nickel alloyed gold ring in your finger when you have allergic difficulty.Why not get on bended knee with a 24kt gold rose in one hand and also a engagement ring during the other and propose to your lover. There's nothing at all wrong with this, it shows genuine appreciate and displays you can take the time and care about your lover. Distinctive colors of your roses have distinctive meanings. The color for adore is red, plus the color for friendship, or falling in love is yellow with red recommendations. Think about if one among these was tipped with 24kt gold? The recipient is positive to say yes. In offering a rose also at a proposal is extremely distinctive. It means for all which might be present and much more the giver at the same time. The giver can delight during the encounter of their enjoy and swim during the happiness of each.It's a known fact the look for the proper form of card can never be uncomplicated. One among the ideal items to complete, to cut back the time taken to look for the proper one particular, should be to know beforehand what you are truly searching for. For those who go out on the industry, you'll discover a quantity of shops which promote this kind of items. On the other hand, except if you understand everything you want, you could possibly effectively shell out the complete day and not discover the proper merchandise to present. It's superior to know that you can find several varieties of Christian greeting cards offered while in the marketplace. Just ensure that you have an idea of what you are trying to find just before you decide to shop for it. Based upon the event, you might be planning to gift your family member or pal with one among the Christian greeting cards, you would locate a wide selection of them in numerous stores.The bicycle clothing may possibly includes cycling shorts (fitted shorts, baggy shorts and bib shorts), jerseys, rain gear, base layer clothing, helmets, gloves, sneakers, socks and headgear. Cycling shorts is usually during the form of fitted shorts, baggy shorts and bib shorts and built to improve the comfort degree and efficiency whilst cycling. These shorts are typically created from nylon spandex fabric which has terrific stretch and durability. Spandex can also be mixed with other yarns this kind of as polyester which presents breathability and protects the skin of your legs from repetitive friction towards the bicycle seat or frame. These shorts assistance in evaporating the moisture from your skin to prevent rashin on the skin and minimizes wind resistance by increasing aerodynamic efficiency. Commonly, these shorts possess a padded stretch liner created within which helps in wicking moisture and stopping chafing. The pads are largely produced from several layers of polar fleece and can differ in thickness. Fitted biking shorts are manufactured in four panels, 6 panels and 8 panels and permit the grament to get contoured to fit your body in the cyling place. The a lot more the quantity of panels, the extra curvature is often allowed to the physique. Bib shorts are held by braces or suspenders as an alternative to elastic waistbands.
These watches not only portray your design but additionally include an x-factor for your persona. The first of the Cartier watches was made from the 12 months 1904 by Louis Cartier for his Brazilian aviator friend, Alberto Santos-Dumont. This view, named as Santos, marked the beginning of the new brand during the market. The fashion, substantial precision, top quality and brilliantly made straps and watchcases created these watches one among the most desired ones amid various sections of society trying to find superlative top quality timepieces. Every model introduced by the brand; be it a quartz movement or mechanical movement 1, exhibits a perfect blend of time-ordered traditions and present day technologies. You may also decide on from watches with guide or automated winding mechanisms, whichever you will be comfortable with. However, every variant would enrich your type and class in its own way. Some of probably the most talked about Cartier models contain Santos de Cartier, Ronde Louis Cartier, Need to de Cartier as well as the sporty Roadster.cartier love bracelet yellow gold
view 301.CI.8710.NR professional or even a timepiece 301.PI.500.RX connoisseur will 301.PX.1180.GR come across it definitely is hard to 301.SM.100.RX.CVL07 figure out if 301.GI.5290.RG 301.PX.130.RX.114 a timepiece is an imitation 2841-36b30-64b 278280-2004 or 301.AM.130.RX.YCM07 not. 301.PE.2180.RW.1104 279/80 Although 301.PX.1180.RX 301.Computer.3180.GR 301.PX.130.RX.174 these Chopard 272.three.36 imitation 301.PE.2180.RW versions visually resemble to 301.CT.130.RX such 301.EI.5190.RB 301.PI.500.RX.114 a degree 301.PX.130.RX the real 27/7471-5002 301.CB.1001.RX timepieces, tips on how to 277.931.91.0371 make certain 279.81.70 the view that you invested 301.CI.1190.GR.ABB09 quite 301.CX.134.RX 301.CX.130.RX penny 301.CM.131.RX.LUN06 on 301.CM.131.RX.LUN06 is the authentic? Generally, it could be 2SWASGMT.S01A.K06B having said that feasible to 2925-3430-53B 301.CI.1770.RX inform the distinction in the event you ever do amazing preparation ahead 301.Pc.1007.RX.094 of buying 28/8914 them. An 2900.51.82 added 300.14.20 lead to why a lot of on-line watch purchasers search for TAG Heuer brand identify watches is thanks 301.CB.1001.RX for the distinctive types which are obtainable. Each men and females will 2852HS acquire which the 301.CX.130.RX kinds 277-933-06-V793-AB12 using the watches produced by TAG Heuer are unique and high-class in nature. 288347-3012 In place of other view rivals, TAG 270.3.36 watches are 301.CK.1140.RX likely to exude slightly significantly more luxury and a smaller way more uniqueness than other brand names. People who 2SWAS.B13A.K06B are buying 277-931-06-0371-AN12 for watches 2CRBS.B01A.L31B will receive that their lookup tends to target TAG Heuer brand identify watches resulting 277000-0007 from your total seem 27/7471-5001 and really feel from the luxury view. An fascinating truth you 301.CE.1201.RX may or could possibly not know: Globally Watch Firm (IWC) is quite possibly 277.931.91.0371 the 301.AI.460.RX.114 one particular primary 301.AI.460.RX.194 Swiss observe company 3006-200 whose founder was an American! Nevertheless, this is wherever it becomes a little bit additional tough and wherever your odds of picking up a fake pair are largely heightened. It is actually vital, for that reason, for making absolutely sure which you spend shut consideration on the description and constantly examine any pictures from the Glasses as closely as you possibly can. Any description that subtly hints at the Glasses currently being counterfeit really should level you in an additional course. One example is, frequently you might find descriptions using words like 'inspired by', or 'like', and hence you ought to ensure that you steer clear of them. When you have essentially made your purchase a clear inspection is doable and also you really should always ensure that you have obtained from a vendor who features a total dollars back guarantee should really you not be content together with the quality of the item. In fact simply just giving a complete refund is generally a typical indication the objects are real. Any time you do get your hands on the solution make certain you test for your excellent on the design and the components which have been made use of. In short, acquiring this view is as superior as getting the view of your year! Rolex Daytona and Replica Swiss Daytona are again two different watches and hence it differs on the grounds of price and material on the watch. Moreover when you are shopping for the view it truly is must to seem more than the characteristics of your model to know your choice and need.
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Top Teams form the country were invited to participate in the
first Premier Cup, a prize money tournament which teams are invited
by order of their rankings. 8 teams converged on Four Bridges
CC to compete for $5,000.00 in prize money put up by Premier
Platform Tennis.
final results
elite event of Platform Tennis concluded on Sunday
w/ one of the sports most recognized names holding
the trophy as the 1st ever Premier Cup Champion. Flip
Goodspeed(Grand Rapids)/Scott Mansager(Chicago) def.
Bill Fiedler(Chicago)/Scott Estes(Toledo) 6-1, 6-3. "This
type of event brings a whole new element of excitement
to our great sport" says Tournament Director Jason
Gray and Dir. of Racquet Sports @ Four Bridges C.C. The
Premier Cup has already gained national recognition from
other areas of the country as clubs are interested in
hosting the sports premier tournament next year!
-Jason Gray
The format was as follows:
- Two brackets of four teams each
- Bracket #1
- Goodspeed/Mansager
- Gambino/Ohlmuller
- Estes/Fiedler
- Bennet/Pierce
- Bracket #2
- Anderson/Stulac
- Cochrane/Marino
- Caldwell/Cordish
- Jagodzinski/Mott
Day one: Starts with a round robin format with each team playing each of the
other 3 in a 10-game pro-set. The two teams with the most wins in each bracket
advance to Sunday's semifinals.


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Ratege Early Origins
Durham, where they held a family seat from early times and their first records appeared on the census rolls taken by the ancient Kings of Britain to determine the rate of taxation of their subjects.
Ratege Spelling Variations
Ratege Spelling Variations
Spelling variations of this family name include: Reddoch, Reddock, Redick, Reddish, Raddish and others.
Ratege Early History
Ratege Early History
This web page shows only a small excerpt of our Ratege research. Another 275 words (20 lines of text) covering the years 1490 and 1597 are included under the topic Early Ratege History in all our PDF Extended History products and printed products wherever possible.
Ratege Early Notables (pre 1700)
Ratege Early Notables (pre 1700)
More information is included under the topic Early Ratege Notables in all our PDF Extended History products and printed products wherever possible.
The Great Migration
The Great Migration
Some of the first settlers of this family name or some of its variants were: John Reddish who settled in Virginia in 1623; William Reddish settled in St. Christopher in 1635; Alexander Reddock settled in Virginia in 1653; John Reddock settled in Virginia in 1650.
Ratege Family Crest Products
Ratege Family Crest Products
See Also
See Also
Other References
The Ratege Family Crest was acquired from the Houseofnames.com archives. The Ratege Family Crest was drawn according to heraldic standards based on published blazons. We generally include the oldest published family crest once associated with each surname.
This page was last modified on 18 November 2013 at 15:46.
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KMC Laboratory Technician Recruitment 2017 for total number of 100 Vacancies application notification are given below. Read more about Syllabus, Admit Card in below passage.
KMC Laboratory Technician Recruitment 2017:-
KMC Laboratory Technician Recruitment 2017:-As per the official recruitment notification there are total number of 100 post are vacant in laboratory technician in the department of Kolkata City NUHM Society. All the candidates are eligible who fulfill the eligibility criteria ( Higher Secondary pass from a recognized board and Diploma in Medical Laboratory Technology) and apply for the post before last date 30 March 2017. The board soon conduct examination (Practicle Test) on different allotted centers on due dates. The submission of application form process is offline and candidates must send all required documents to the particular address. Its a good opportunities for all those who are medical student and willing to do job in the particular field. Its important for all those candidates who wish to apply for the post to check all necessary stuff in the official advertisement and direct link of advertisement is given in the below passage. Apply soon KMC Laboratory Technician Recruitment 2017.
KMC Laboratory Technician Admit card 2017:-
|Important Events
|Important Dates
|Official Board
|Kolkata City NUHM Society
|Application Mode
|Online Application
|Last date of application form
|Starting date of application form
|Official website
|Exam date
|Soon declared
KMC Job Details :
- Post Name : Medical Officer (Part Time)
- No. of Vacancy : 104 Posts
- Pay Scale : Rs. 24000/- (Per Month)
Eligibility Criteria for KMC Recruitment :
- Educational Qualification : MBBS from a MCI Recognized Institute with 1 year compulsory internship .
- Nationality : Indian
Age Limit : Maximum age limit is 66 years (as on 01.01.2017)
Job Location : Kolkata (West Bengal)
Selection Process : Selection will be made on through Interview.
Application fee : There is no Application Fee.
How to Apply KMC Vacancy : Interested candidates may appear for Interview along with Prescribed Application form & attested copies of certificates & Document at the time of Interview.
Venue of Interview : Room No-254, 2nd floor, PMU Kolkata City NUHM Society, 5 S.N Banarjee Road, Kolkata-700013.
Important Dates to Remember :
- Date of Interview : 08.03.2017 at 11.30 AM
Important Links :
For more related recruitment is to be given in the official website https://jobsinfo.in/. Check all those related information and get job.


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To those who think no one is harming Christians in Egypt.... UPDATE eyewitness accounts
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[quote:Riker:MV8xMzQ1NDMzXzIyMTYyNzUxXzQ0OEQzNzhE] [quote:Anonymous Coward 1244935] peace and protection :hf: hope you are ready for the storm Riker! We are supposed to get all ice here in Maryland. Won't be going to see Punx. Phil this year! [/quote] Thank you !!! :hf: It hasn't started here yet but it's supposed to soon. Right on time too; my bday is Thursday and it's always cold and blustery weather :-) [/quote]
Original Message
I'm so freaking sick of people saying that Christians and Muslims are equally violent. Not all Muslims are violent and not all Christians are the epitome of peace....
But I am getting emails from friends with family and/or missionaries in Egypt and it is BAD for Christians in Egypt right now, worse than usual. They can't get out. As if the bombing of churches wasn't enough, now they are fearing for their lives with the Muslim uprising.
I don't care if this is a "movement of the people" if what the people want is to harm the minority!!!
Please pray for peace and protection.
Received an update today:
Eye-Witness Account from Egypt
Dear Friends,
Egypt is finding its way towards more freedom, but please don't believe what the international media is saying! There is a plot to topple the president, and Egyptians do not want that. I have been on the streets with many hundreds of thousands, probably even millions, who want stability, dialogue, and change with Mubarak until the elections in September. There are forces taking advantage of the situation and twisting facts and figures to portray to the international community a distorted image to topple the government. The vast majority of people in the streets in peaceful demonstrations (I speak as an eyewitness because I took to the streets myself) are carrying banners saying things like:
- yes to stability, yes to Mubarak
- give change a chance
- we are sorry Mr. president
- we accept dialogue, we trust you
- no to El-Baradei, no to the muslim brotherhood (many like this one)
- we are the Egyptians, where is Al-Jazeera, let them come and see
- no to corruption, no to vandalism
- we got what we asked the president for, so why are people still in Tahrir? Who are they? What do they want? etc., etc., etc.
We met with people who were in the original protest in Tahrir square who decided to join us saying: we got what we asked for, and now we accept Mubarak's changes and proposals. I left the street with my wife around 4:15 pm, and the numbers in the neighbourhood of Mohandessin where the crowds were gathered had swelled to easily over a million. As we drove home we saw the same slogans on banners all over the city, on cars, on walls, on shop windows. We learned that similar demonstrations are taking place ALL OVER THE COUNTRY, IN MANY DIFFERENT CITIES. THIS IS THE CRY OF THE PEOPLE OF EGYPT THAT IS BEING TOTALLY IGNORED BY THE INTERNATIONAL NEWS MEDIA. Is this on purpose??!!! I am perplexed!!!
I am wondering: How come CNN, the BBC, and others are reporting ONLY the anti-government protests as the voice of the people? This is not JUSTICE, this is not the TRUTH. There have been reports that these people are being paid by the government. NOT TRUE! I was there with many, many others. I SAW THE STREETS. Now to the situation in Tahrir square. Only a few people (hundreds?) are still there from the original protesters. They have been slowly replaced by other HIGHLY ORGANIZED GROUPS. They all have the same model of cell phones. They all have the same blankets (eye witnesses). THESE ARE NOT THE PEOPLE OF EGYPT.
Some witnesses claim that they don't look Egyptians, and don't sound Egyptians (different accent, different dialect). THIS IS A BIG ORGANIZED COUP TO TRY TO CONVINCE THE WORLD THROUGH THE MEDIA THAT EGYPT WANTS MUBARAK TO GO, AND THE MEDIA IS PART OF THE DECEPTION. People in Tahrir square are escalating the situation on purpose to topple President Mubarak FOR THEIR OWN HIDDEN AGENDAS. This is TYPICAL OF THE MUSLIM BROTHERS, AND EVERYBODY IN THE STREETS OF CAIRO KNOWS THIS. We heard people on the streets saying that the plot to take over the country is now clear. THE INTERNATIONAL MEDIA DOESN'T WANT YOU TO KNOW THIS.
The escalation of violence in Tahrir square is because of this. Egyptians who love Egypt, the millions that took to the streets yesterday, want this to end. They fully understand that president Mubarak is between a rock and a hard place, that he cannot quench the unrest in Tahrir through the army, so the people want to go to Tahrir to disperse the crowds there by themselves. People in Tahrir are vastly outnumbered. If Egyptians go the Tahrir square to take control of the situation, more chaos will erupt, giving a chance to the international media to blame the President even more.
Don't believe an opportunist like Mohammed El-Baradei who is deceiving the international community into putting pressure on president Mubarak to leave immediately so they can fulfill their own agendas. Franklin Roosevelt said, "Nothing happens by accident in politics. If it happened, somebody made it happen." And this is NOT what Egyptians want! I am an Egyptian. My wife and I were on the streets on Tuesday and Wednesday in support of the current regime, and we saw millions on the streets. If Egypt falls, then neighbouring countries are going to fall one after the other. Someone is behind this!
Please help us uncover the deception and spread the truth. Please share with everybody.
Dear Intercessors,
We have been following the events carefully as they unfold in Egypt and across the Middle East. As usual, things aren't always as they seem in political circumstances. Below you'll find an email that I received from a witness on the ground in Egypt. This is a grave situation. Leaders from Egypt have been calling and beseeching us for intercession for their nation. From their perspective, the Christians are in grave danger if Mubarak leaves, as they feel he has kept the radical extremists at bay.
Here are some prayer points:
1. Pray that God will reveal every secret plot the Muslim Brotherhood has in Egypt.
2. Pray that every scheme that is Al Quaeda backed will be exposed in all levels of Egyptian culture, including the police.
3. Pray for protection of Christians across the Middle East
4. Pray for a smooth transition to a secular government.
5. Pray that the military of Egypt will not stand with pro-Muslim Brotherhood factions.
6. Pray that factions will not blow up the oil pipeline and that the Suez Canal will stay open for international commerce.
7. Pray that this Islamist revolution will not be successful and that they will not be able to establish an Islamic Caliphate.
8. Pray for protection for Israel from the terrorists that would sneak across the border into Gaza while the borders are unguarded.
9. Pray for comfort and peace for those who have been caught up in the midst of their nations being destabilized.
10. Pray for a movement of unified fasting and prayer to sweep across the believers in Jesus and powerfully impact the Middle East.
11. Pray for an Isaiah 19 highway of holiness to be established.
12. Pray for a Great Awakening to take place, with thousands of souls swept into the Kingdom of God.
Standing in the gap!
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Brooklyn, NY, US
Architecture, urbanism and environment are my passions, I strive for tactile results and thrive in either a collaborative or solo atmosphere. A nuanced understanding of the urban condition today and historically is vital and always informs my approach to problem solving. Each city is different, but many good cities share the same underlying principles. How can we maintain the character and individuality of cities (and thus structures) while simultaneously bringing them into the 21st century? I believe the answers are rooted in sustainability and community. Within those two realms are a wealth of resources that can help us as designers to create not just best practice, but innovative practice.
**I am currently logging my IDP hours and actively working towards becoming a licensed architect.**
Parsons The New School for Design, New York, NY, US, Part-Time Faculty
Perez, APC, Brooklyn, NY, US, Architectural Designer - Freelance
Perez, APC, Brooklyn, NY., Architectural Designer
Indamine Architecture, Brooklyn, NY., Designer
Pierre Frey, New York, NY, US, Freelance Drafter
Passive House Institute US (PHIUS), New York, NY, US, Coordinator_Training Event
AHDLI Symposium Summer 2012 NYC, New York, NY, US, Design Resource Assistant
Indamine Architecture, Brooklyn, NY, US, Assistant to Leah Kreger, RA.
Droog, New York, NY, US, Sales & Visual Merchandiser
Parsons The New School for Design, New York, NY, US, MArch, Architecture
As a M.Arch student much of my focus involved sustainable, green and resilient design. I've had the opportunity to design academically, a Passive House row home for Habitat for Humanity Philadelphia, a public library in the East Village, mixed use affordable housing in the Bronx, an alternative to the FDR highway that would protect that Manhattan shoreline from rising waters, and a recycling and marine transfer station on the East River.
The New School for Social Research, New York, NY, US, Bachelors, Urban Studies
My initial studies involved fine arts and photography, and eventually focused on urban studies, architecture and architectural theory.
Pratt Institute, Brooklyn, NY, US, BArch, Architecture
I attended Pratt School of Architecture for one year, before deciding to return to The New School to finish my undergraduate studies, with the intention of pursing architecture again in Graduate school.


text: This 4-star-superior hotel in Eschborn offers an outdoor pool, air-conditioned rooms, and great links with the A5 motorway. It is just 7 km to Frankfurt`s city centre and the International trade fair grounds. The Mercure Hotel Frankfurt Eschborn Ost has comfortable, air-conditioned rooms with work areas, satellite TV, and views of the Frankfurt skyline. Guests can also relax under palm trees in the Mercure`s poolside beach area with bar and club. Regional and international dishes are served on the terrace in the Big Ebbel restaurant. Drinks are served in the Skyline Bar. Private parking is available at the Mercure Frankfurt Eschborn Ost.


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by Wiley Clapp - Friday, August 21, 2009
The 356 TSW was an interesting cartridge that lasted only very briefly and is now out of print. It stands for 356 Team Smith & Wesson and was offered in several S&W autos as well as a few Centennial revolvers.
The S&W engineers created the little 9 mm round for the early 1990s IPSC venues. Their goal was to use existing pistol platforms to build competitive autos.
Loaded to the same overall length as the 9 mm Luger, the 356 TSW drove 147-grain JHPs fast enough to do the trick. Using a strong case about 21.5 mms in length, the 356 TSW made major with room to spare. At one point, I had several guns for evaluation and enjoyed shooting them very much. Basically, it was a way to hot rod the 9 mm round and it worked.
For one reason or another, the round was barred from competition and passed from the scene. S&W may still make some of the guns for sales in Europe, where things are different.
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Fujifilm announced the Instax Share SP-2, which allows you to print digital photos onto instant film.
The Instax Share SP-2 is the successor to the SP-1 and it prints quicker and it's quieter when doing so. The pocket-sized device prints onto Instax's mini format instant Instax film, allowing you to keep your digital photos in physical form.
There's a free Share app that connects your smart device to the Share SP-2. It also lets you print photos side-by-side or four in one slide as part of a collage, and lets you apply filters. You can even add info like the place, date, and weather to an image.
The Instax Share SP-2 will be released in the US in mid-July. Fujifilm is yet to announce release dates elsewhere.


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Considering it was his first public interview in 10 years, Rahul Gandhi was understandably nervous. Things started badly for the Indian Congress leader, got worse towards the middle and the less said about the end the better.
Agreeing to answer prominent journalist Arnab Goswani's questions "specifically", the Indian National Congress leader issued a disclaimer that he would take time to come to his point. However, in most cases, he never really got there.
It was evident that the "shehzada" (prince), as he is called by his chief political rival Narendra Modi, was ducking and diving to avoid certain questions. Nothing wrong with that - after all most global political leaders have honed the skill of finding their way around probing queries from those pesky journalists. Gandhi, on the other hand, has a lot to learn.
He seemed completely unprepared for Goswani's questions and started shuffling about in his chair like a nervous schoolboy when asked whether he was afraid of going up against Modi for prime minister.
Rahul Gandhi has sincere naivete.I would definitely vote for him as house captain in my school. — Naomi Datta (@nowme_datta)
January 27, 2014
Negative: Rahul got stuffed a lost the little respect people had Positive: Mummy probably going to buy him a Playstation 4 to cheer him up — Khamba (@gkhamba)
January 27, 2014
It was downhill from there.
It's not as though he wasn't saying the right things. The only problem was that Gandhi's replies were not even remotely related to Goswani's questions. Yes, the women of India need to be liberated and empowered, but how is that relevant to the involvement of Congress leaders in the Anti-Sikh riots of 1984?
When further prodded by the journalist, the Congress leader finally touched upon the sensitive topic, but showed he was a little disconnected from reality.
"All I'm saying is there is a difference between the 1984 riots and the Gujarat riots. The difference is that the government of the day in 1984 was not aiding and abetting the riots. That is all I'm saying."
Perhaps the palace messenger forgot to inform the "shehzada" that Congress leaders were actively involved in abetting the chain of events.
"When things got hairy, he spoke about [the] system and empowerment," said analyst Siddharth Varadarajan.
In fact, Gandhi liked the word 'system' so much that he mentioned it 72 times during the course of his one-hour-twenty-minute-long interview.
On the other hand, his nemesis Narendra Modi seemed to be the furthest thing from his mind and was mentioned just three times. Rather strange considering that a number of the Times Now anchor's questions were focused on Modi. When asked about the prospect of a one-on-one debate with the BJP leader, Gandhi again chose to take refuge in the 'system' he wishes to 'empower'.
The sound you hear is fans of the BJP coming in their pants. — Overrated Outcast (@over_rated)
January 27, 2014
There were other questions raised by Goswani over corruption within the Congress Party, dynastic politics in India and Rahul's supposedly fragile character; all of which the politician side-stepped. His interview started sounding like a well-versed speech about his determination to change the system and he never effectively answered a single one of the probing queries.
If Rahul Gandhi wishes to change the face of Indian politics and empower his people, he will first and foremost have to be taken seriously. Gauging from the reaction of social media and other analysts, the day when he can become a force to be reckoned with is a long way ahead.


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About Pork Shoulder Steak Edit
Pork shoulder arm steak has the same muscle and bone structure as pork shoulder arm roast, only cut thinner. It contains round arm bone and the meaty part of the arm picnic. The outside is covered with a thin layer of fat. It is usually prepared by braising or pan-frying.
Pork shoulder blade steak is cut from pork shoulder blade Boston roast. It contains blade bone and several muscles and is usually prepared by braising, broiling, grilling, pan-broiling, or pan-frying.


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Drew Barrymore & Will Kopelman Wedding Day Kiss
Drew Barrymore and her new husband Will Kopelman share a wedding day kiss!
The newly married couple were spotted kissing as they left their wedding in Montecito, California in a classic Mercedes. The star studded wedding included several high profile guests including Cameron Diaz, Reese Witherspoon, Jimmy Fallon and Busy Philips. It's also been rumored that Drew is expecting her first child but wanted to be married before announcing the news.
Check out Drew Barrymore and Will Kopelman on their wedding day! (Fame/Flynet)


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Extract/Partial Mash Brewing / Re: ALMOST perfect brew day (question)« on: November 15, 2014, 01:05:15 PM »
Yep HopShot residue. No issues with that in the beers I have used them in.
This section allows you to view all posts made by this member. Note that you can only see posts made in areas you currently have access to.
That is very cool.I am in downtown Waukesha at Bernie's taproom. http://berniestaproom.com/
Quick question for Thirsty_Monk. I will be in Southern WI and IL for a few days this upcoming week. Do you distribute that far south? Would love to try your pilsner.
I do not have any package products down there.


text: In October 2015, Oklahoma was awarded a $225,000 grant from the C.S. Mott Foundation to create a statewide partnership focused on Out of School Time programs. The Oklahoma Partnership for Expanded Learning (OPEL) was created to support Out of School Time programs to coordinate a unified vision to inform policy and quality afterschool initiatives.


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Wyoming (Cassette)
WYOMING! 1838. The first wagon train to Oregon country moves slowly across the Great Plains, reaching the foothills of the Rocky Mountains in early autumn. Led by wagon scout Whip Holt, the five-hundred-strong community of courageous men, women, and children prepare for the bitter winter ahead. Among them are dreamers and drifters, foreigners and natives, friends and enemies. But it is a pair of strong-willed women who are Whip s most unexpected allies: the outspoken Cathy van Ayl, widowed and alone at twenty-four, and the beautiful half-Arapaho, half-Cherokee La-ena, who was once Whip s mistress. Before these brave pioneers lie endless miles of hardship and danger, treachery and death. Beneath their struggle for survival lie hidden rivalries, unspoken secrets, and burning jealousies that threaten to consume them all. But in their hearts beats the dream of a new beginning in a promised land the unstoppable dream of America itself.
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text: Pronounced Broo-NELL-oh. A grape varietal, used in the production of powerful red wine, produced in the Tuscany region of Italy. Brunello, referred to as "little dark one", is actually a strain of Sangiovese Grosso. Brunello is the only grape varietal in the costly and rare Tuscan Red, Brunello di Montalcino D.O.C.G. wines.