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Axon regeneration is a medically relevant process that can repair damaged neurons. This review describes current progress in understanding axon regeneration in the model organism Caenorhabditis elegans. Factors that regulate axon regeneration in C. elegans have broadly similar roles in vertebrate neurons. This means that using C. elegans as a tool to leverage discovery is a legitimate strategy for identifying conserved mechanisms of axon regeneration.
Caenorhabditis elegans is a comparatively new model for the study of axon regeneration. Here, we review progress in the field since its beginning in 2004 and discuss directions that future investigations might follow.
Damaged axons can in some cases regenerate, restoring function to nervous systems after injury or disease. Regeneration of injured neurons is thought to be initiated by signals arising from the injury site (20, 66, 74). These injury signals promote remodeling of the cytoskeleton and plasma membrane at the site of injury, resulting in the generation of a new growth cone. Regulated changes in gene transcription and local and somatic protein synthesis are also required for successful axon extension during regeneration.
The response of a specific neuron to injury is the result of the interplay between various pathways that promote or suppress regeneration. In part, regeneration is regulated by signals from the neuronal environment. For example, myelin-derived signals [Nogo, MAG (myelin-associated glycoprotein), and OMgp (oligodendrocyte myelin glycoprotein)] and chondroitin-sulfate proteoglycans (CSPGs) inhibit regeneration in the vertebrate central nervous system (CNS) (97). However, regeneration is also regulated by intrinsic factors, and altering the intrinsic state of a neuron can improve regeneration, even in the presence of inhibitory factors (58, 59, 65). Thus, building a complete model of the mechanisms that regulate and mediate regeneration is critical to understanding how regeneration works and why it often fails.
The nematode worm C. elegans has recently emerged as a model for axon regeneration. Axons can be severed through the transparent skin of the worm using a pulsed laser, triggering regeneration (96). Regeneration can then be studied in vivo at single-axon resolution (Figure 1). When this technique is combined with the powerful genetics and other tools available in C. elegans, it becomes possible to investigate the complex genetics and cell biology of axon regeneration more rapidly and in greater molecular detail than previously possible (27–29, 36, 57, 92, 95). An important limitation of C. elegans for these studies is the relative simplicity of the extrinsic neuronal environment. Worms have no myelin and no invading macrophages. Thus, the worm is a poor model for certain questions regarding the interaction between an injured neuron and its environment. In many cases, however, the simple environment surrounding C. elegans neurons can accelerate discovery by removing confounding factors.
An essential question of any biological model is its relevance to human biology. A precise comparison between the mechanisms of axon regeneration in C. elegans and those in humans (or at least vertebrate models) is not yet possible because these mechanisms are incompletely understood. However, results so far are encouraging: (a) In C. elegans, as in vertebrates, cAMP (cyclic adenosine monophosphate) acts to promote axon regeneration after injury (8, 28, 58); (b) the DLK-1 mitogen-activated protein kinase kinase kinase (MAPKKK) promotes regeneration in both C. elegans and mice (36, 43, 95); and (c) axon guidance molecules are important for regeneration in worms and mice (27, 37, 92). Thus, many factors that regulate axon regeneration in C. elegans have broadly similar roles in vertebrate neurons. In this review, we describe how C. elegans is used to study regeneration and summarize recent work that has begun to describe the mechanisms that regulate and execute regeneration.
A major challenge in studying regeneration is that regeneration only occurs only in response to nerve injury. In mammals, injury is delivered by surgically cutting or crushing a nerve or a section of the spinal cord. These techniques are not easily applied to C. elegans because of the small size of the worm. Thus, the study of regeneration in C. elegans has required development of two novel injury models: laser axotomy and the β-spectrin mutant background.
In 2004, it was found that laser pulses can sever GABA (γ-aminobutyric acid) neurons in C. elegans in intact, living animals and that regeneration can be observed after surgery (96). The first laser used to cut axons in C. elegans was a sophisticated, amplified Ti-sapphire laser (96). It is now clear that many varieties of pulsed laser can cut axons and elicit regeneration, including unamplified Ti-sapphire lasers (92), coumarin dye lasers (92), Nd:YAG (neodymium-doped yttrium aluminum garnet) lasers (68), and solidstate lasers (89). In addition to the GABA motor neurons, laser axotomy has been used to sever many other neurons, including the PHA phasmid sensory neurons (92), the PLM and ALM touch neurons (92), the sensory neuron AFD (18), the AWB chemosensory neuron (92), the AVM mechanosensory neuron (27), the DA/DB motor neurons (27), and the HSN neuron (27). There are no examples of neurons that cannot be cut, suggesting that laser axotomy can be used on any neuron in the worm.
Pulsed lasers sever axons by generating free electrons, by causing the formation (and collapse) of nanoscale bubbles, and by generating free plasma that results in a cavitation bubble (10). The precision of laser axotomy enables very accurate and controlled injuries at the single-neuron level. For example, Wu and colleagues (92) found that the success of regeneration in the ALM and PLM neurons depends on how close to the cell body the injury occurs: Injury close to the cell body results in increased regeneration. Similarly, Gabel and colleagues (27) showed that in the DA/DB motor neurons cutting closer than 20 µm to the cell body elicits growth from the cell body rather than the axon stump. A potential limitation of laser axotomy is the requirement to individually target and sever each individual axon that is to be studied. However, diligent application of this technique has yielded nearly all current data, including one study that analyzed hundreds of genotypes (15). One way to increase the speed of laser axotomy is by using a microfluidic platform to immobilize worms, combined with an automated axotomy system (3, 32, 73). A microfluidic and automated axotomy system has been used to screen for chemical modifiers of regeneration (75). Such systems hold the promise of significantly increasing experimental throughput.
β-spectrin is a major and essential component of the membrane skeleton, a dense two-dimensional protein mesh that is tightly associated with the plasma membrane in most metazoan cells (6, 7). In C. elegans, β-spectrin is encoded by the unc-70 gene (34). Neurons in animals lacking unc-70/β-spectrin develop with normal morphology. However, these neurons are fragile, and their axons break spontaneously when the mutant animals hatch and start to crawl (35). Broken axons regenerate, so regeneration can be studied in unc-70/β-spectrin mutant animals without the requirement for experimentally delivering nerve injuries.
Axon breaks in unc-70/β-spectrin mutant animals occur stochastically and may even recur in the same axon (35). Thus, it is more difficult to collect quantitative single-neuron data on regeneration using the unc-70/β-spectrin mutant background than using laser axotomy. However, because regeneration occurs spontaneously and constitutively, it is far easier to collect many animals that are regenerating. The unc-70/β-spectrin mutant background therefore may have advantages over laser surgery for experiments that require very large numbers of animals, such as genetic screens and genomic and proteomic studies. For example, this sensitized mutant background was successfully used in an unbiased RNA interference (RNAi) screen for genes affecting regeneration (36).
An important and unresolved question is the extent to which different injury models—the use of different types of lasers, the study of different neurons, or the spontaneous breaks in unc-70/β-spectrin mutants—converge on common regeneration mechanisms. Detailed studies using femtosecond lasers to sever C. elegans axons show that dramatic changes in regeneration occur as a result of changing the number and energy of laser pulses, even though the total amount of energy applied is constant (10). Even the choice of fluorescent marker can influence regeneration (92). In some cases, such as the dlk-1 mitogen-activated protein kinase (MAPK) pathway, regeneration mechanisms show consistent effects across multiple neuron types, markers, and even injury models (36, 95). In other cases, however, the extent to which mechanisms remain consistent across different cells, markers, and injury models remains to be determined.
Regeneration is regulated by intrinsic factors, and altering the intrinsic state of a neuron can improve regeneration, even in the presence of an inhibitory environment (58, 59, 64). Intrinsic pathways can also inhibit regeneration (17, 63, 81, 84). Both pro- and antiregeneration pathways have been described in C. elegans and are reviewed in this section.
Calcium and cAMP (cyclic adenosine monophosphate) signaling promote axon regeneration in a variety of systems (4, 13, 102). Increased calcium influx as a result of injury can result from entry of extracellular calcium through the breached axonal membrane, activation of voltage-gated calcium channels in response to depolarization, and release from intracellular stores (47, 101). Although increased intracellular calcium can result in neuronal cell death in some cases, calcium signaling can promote membrane resealing and regeneration in surviving neurons (71, 93). Increased intracellular calcium triggers multiple signaling events, including an increase in cAMP through modulation of adenylyl cyclase activity (19).
Calcium and cAMP also promote axon regeneration in C. elegans (28). Axotomy of the PLM touch neuron axons results in an acute increase in intracellular calcium. This increase is at least partially dependent on the voltagegated calcium channel subunit EGL-19 and the inositol triphosphate receptor ITR-1. Blocking the function of EGL-19 or ITR-1 reduces the acute increase in intracellular calcium and also reduces regeneration. Similarly, differences in the acute calcium increase in wild-type animals correlate with differences in the length of axon regeneration. Thus, intracellular calcium entry promotes regeneration in C. elegans. A major effector of calcium in regeneration is the adenylyl cyclase-cAMP-protein kinase A (PKA) signaling cassette. Interestingly, BAPTA (1,2-bis(o-aminophenoxy)ethane N, N, N′, N′ tetraacetic acid) treatment reduces the calcium baseline and reduces regeneration without affecting the size of the calcium transient. This suggests that total levels of calcium, rather than the relative size of the increase, are the final determinant of regeneration.
Increased neuronal excitability results in more intracellular calcium, whereas decreased excitability results in less (reviewed above) (28). Consistent with the finding that calcium mediates regeneration, mutations that decrease excitability result in poor regrowth after axotomy, including the sodium pump nkb-1, the sodium channel unc-8, and the stomatins unc-1 and unc-24. Similarly, mutations that disrupt the synthesis of acetylcholine (cha-1) or acetylcholine transport (unc-17), or that disrupt the deg-3 subunit of the nicotinic acetylcholine receptor, result in decreased regeneration (15). Conversely, genetic lesions that make neurons more excitable, such as loss-of-function mutations affecting the slo-1 K+ channel or the K+ channel regulator mps-1, enhance regrowth (15).
The dlk-1 MAPK regulates regeneration in both GABA motor neurons and sensory neurons (36, 95). An unbiased RNAi screen in unc-70/β-spectrin mutants for genes affecting regeneration identified dlk-1 as a candidate regeneration gene (36). dlk-1 is a MAPKKK that functions in a pathway with the MAP kinase kinase (MAPKK) mkk-4 and the p38 MAPK pmk-3 (56) (Figure 2). A major target of pmk-3 is the MAPKAPK (MAP kinase activated kinase) mak-2, which functions via the bZip-containing protein cebp-1 (95). This entire signaling pathway is required for regeneration. Loss of any component reduces regeneration, whereas activation of the pathway (by overexpression of dlk-1) increases regeneration (36, 95).
The same RNAi screen in unc-70/β-spectrin mutants identified a second MAPKKK, mlk-1, as also having a significant role in regeneration (36). mlk-1 activates the MAPKK mek-1, which acts upstream of the JNK (c-Jun N-terminal kinase)-like kinase kgb-1 (52) (Figure 2). Again, this entire signaling pathway functions in regeneration. Further, the mlk-1 pathway exhibits significant cross talk with the dlk-1 pathway. In addition, the dlk-1 and mlk-1 MAPK pathways share common negative regulators. rpm-1, an E3 ubiquitin ligase that regulates synapse morphology and axon termination (30, 77, 100), mediates the degradation of mlk-1 as well as dlk-1 (56, 60). Hyperactivation of one or both of these pathways in rpm-1 mutants leads to increased regeneration (36). Both the dlk-1 and mlk-1 MAPK pathways are also regulated by the dual phosphatase vhp-1 (60).
In addition to these common regulators, the mlk-1 pathway is regulated by growth factor signaling. svh-1 is homologous to hepatocyte growth factor, macrophage stimulating protein, and plasminogen. svh-1 is a secreted factor that acts via the receptor-type tyrosine kinase svh-2 to activate regeneration via activation of the mlk-1 pathway. Although svh-2 functions in the injured neuron, svh-1 is expressed in the ADL neurons and functions as an extrinsic signal. Thus, the response of injured neurons is regulated in part by secreted factors deriving from other, uninjured neurons.
How does MAPK signaling facilitate regeneration? Activation of the dlk-1 pathway results in the stabilization of the cebp-1 messenger RNA (mRNA) through its 3′ untranslated region. Some mammalian MAPKAPKs (homologous to mak-2) can phosphorylate mRNA binding proteins, which may result in differential regulation of mRNA turnover (40). Thus, mak-2 may mediate cebp-1 mRNA stability by modifying RNA-binding proteins in response to axotomy. In the end, via the dlk-1 and mlk-1 pathways, axotomy results in localized translation of cebp-1 and also activation of kgb-1. However, the downstream effectors of cebp-1 are not known, nor are the effectors of kgb-1.
Microtubules play a crucial role in axon regeneration. Destabilization of microtubules with nocodazole in regenerating rat sciatic nerve inhibits growth and transforms growth cones into retraction bulbs. Conversely, chemical stabilization of microtubules in the rat spinal cord by taxol treatment reduces the formation of retraction bulbs and enhances growth-cone formation. Retraction bulbs exhibit a disorganized microtubule structure and are static, whereas growth cones show a characteristic bundling of microtubules essential for growth-cone movement and axonal regrowth (22). Manipulation of microtubules results in enhanced regeneration in the mammalian CNS and may present novel therapeutic possibilities (39, 78, 82).
Microtubule dynamics are also important for C. elegans axon regeneration. efa-6 mutations result in increased PLM regrowth after axotomy (15). EFA-6 contains a variable N-terminal domain, a pleckstrin homology (PH) domain, and a Sec7 domain, which has a guanine exchange factor activity for ADPribosylation factor GTPases (26, 61). However, neither the PH domain nor the Sec7 domain of EFA-6 is required for inhibition of regeneration. Rather, the N terminus of EFA-6 inhibits regeneration by regulating microtubule organization. Although loss of efa-6 results in increased regeneration, overexpressing the N terminus of EFA-6 results in dysmorphic growth cones, less regeneration, and a reduction in dynamic microtubules (15). Stabilizing microtubules with taxol suppresses the inhibition of regeneration resulting from EFA-6 overexpression. This suggests that EFA-6 destabilizes microtubules in regenerating PLM axons and leads to failure of regrowth.
The lin-12 Notch receptor is a potent inhibitor of regeneration in C. elegans motor neurons (21). lin-12/Notch null animals regenerate significantly better than wild-type animals, whereas lin-12 gain-of-function alleles regenerate worse. Notch functions in the injured neuron to inhibit regeneration. Further, Notch activation after injury is necessary for Notch to inhibit regeneration: Blocking Notch activation with the γ-secretase inhibitor DAPT (N-[(3, 5-difluorophenyl)acetyl]-Lalanyl-2-phenyl]glycine-1,1-dimethylehyl ester) increases regeneration, even when injury has already occurred.
Notch activation typically involves sequential cleavage of Notch, first by a metalloprotease (site 2 cleavage) and then by the γ-secretase complex (site 3 cleavage). These cleavages release the Notch intracellular domain (NICD) into the cytoplasm. Notch inhibition of regeneration proceeds by this same activation mechanism: Both the sup-17/ADAM10 metalloprotease and presenilin (the catalytic component of γ-secretase) are required for Notch to inhibit regeneration, and NICD overexpression is sufficient to inhibit regeneration.
The activation mechanism for Notch inhibition of regeneration is not known. DSL (Delta/Serrate/LAG-2)-family Notch ligands are single-pass transmembrane proteins (87, 98). In addition to transmembrane DSL ligands, C. elegans expresses multiple secreted ligands thought to activate the receptor in cooperation with transmembrane ligands (16, 25, 45). However, loss of any single ligand does not improve regeneration. One possibility is that multiple ligands can redundantly activate Notch to inhibit regeneration. Alternatively, Notch activation might be ligand independent. In Drosophila, endocytosis of the Notch receptor can be mediated through Deltex in a ligand-independent manner and stabilized by Hif-α (41, 42, 55, 88, 94). Once endocytosed, Notch is cleaved by presenilin, and the NICD can be released into the cytoplasm. Calcium-dependent, ligand-independent Notch activation has also been described (67).
Notch activation is required within a short time frame (2 h) of axotomy to inhibit regeneration. This limited window of activation suggests that Notch may be directly activated by injury and that it signals immediately to inhibit regeneration. This immediate activation would allow enough time for an inhibitor of regeneration to be transcribed and translated to inhibit growth-cone initiation. Alternatively, Notch may be active at all times but may need to interact with other pathways that respond directly to injury or may need to be active continuously because its effectors are short lived. How Notch signaling affects regeneration remains an open question. One possibility that is consistent with our findings is that Notch could act by activating transcription of a direct inhibitor of regeneration. Some targets of Notch signaling have been identified, but their role in regeneration has not been examined.
Synaptic vesicle exocytosis at nerve terminals results in the transfer of vesicle proteins and lipids to the plasma membrane; these are retrieved by endocytosis. Endocytosis— but not exocytosis—is also required for regeneration (15). Mutations in any of three key endocytosis genes (unc-26/synaptojanin, unc-57/endophilin, and unc-41/stonin) result in decreased regeneration. The requirement for synaptic vesicle endocytosis (but not exocytosis) genes in regeneration suggests that regeneration does not depend on synaptic vesicle cycling but on different cellular process. One potential mechanism is MAPK signaling, as the requirement for unc-57/endophilin during PLM regeneration can be bypassed by increased DLK-1 expression (15).
MicroRNAs function in many aspects of neural development and disease (46). Studies in cultured neurons, mice, and zebrafish suggest a role for microRNAs in central and peripheral regeneration (83, 91, 99). Because microRNAs are generally involved in the downregulation of their target genes, a regeneration-promoting role of microRNAs suggests an important role for the negative regulation of inhibitors of regeneration. In C. elegans, the Argonaute homolog alg-1 is required for PLM axon regeneration (15). alg-1 functions in the biogenesis of microRNAs (31), suggesting a role for microR-NAs in regulating regeneration in C. elegans. Although a general regeneration-promoting role for microRNAs has recently emerged, further study of the roles of individual microRNAs may reveal specific roles.
Regenerating axons navigate a very different environment than the one they encounter during development. Although some developmental growth and guidance cues may be maintained in adults, others are lost or replaced by inhibitory factors. Numerous studies in mammalian systems show that extrinsic factors from the glial scar and myelin act as potent inhibitors of regeneration in the CNS (reviewed in 11, 24, 50, 79, 97). C. elegans axons are not myelinated and navigate a very different extracellular environment than mammalian neurons. However, recent evidence suggests that axon guidance factors and elements of the C. elegans extracellular matrix (ECM) also play a role in the regulation of axon regeneration in worms.
Regeneration in C. elegans is often characterized by guidance errors, premature termination, or branching (27, 92). By contrast, development of the nervous system is largely invariant. One mechanistic difference between regeneration and development is the function of axon guidance pathways. During development, ventral migration of the ALM mechanosensory neuron depends on the netrin and the Slit/Robo guidance pathways (27). These pathways depend on extracellular ligands (netrin and Slit) and their neuronal receptors (the netrin receptors unc-5 and unc-40 and the Slit receptor Robo). During regeneration, netrin and Slit also help to guide ALM ventral migration: Migration is defective in mutants for these genes. However, the neuronal receptors are dispensable for this process (27). Thus, axon guidance signals function differently in ALM during regeneration than during development.
A change in response to axon guidance molecules is also found in the PLM neurons. In these neurons, Slit/Robo signaling promotes outgrowth (49). During regeneration, however, Slit/Robo signaling inhibits axon extension (15). Further, accurate regeneration in the PLM neuron in adult animals is inhibited by the vab-1 Eph receptor (92). However, this function is independent of the kinase activity of vab-1 (92), whereas kinase activity is required for vab-1's developmental function (53). Thus, differences in how axon guidance pathways function may account for some of the pathfinding errors during regeneration.
Many axons in C. elegans migrate along the basement membrane that runs between the body wall muscles and the epidermis. This basement membrane is required to maintain attachment of the muscle to the epidermis. Three genes—pxn-2, spon-1, and vab-19—that inhibit regeneration encode proteins that function to maintain muscle attachment. PXN-2 is a peroxidasin, which is a secreted protein that contains a catalytic peroxidase domain, immunoglobulin domains, and leucine-rich repeats. Peroxidases can inactivate metalloproteases, cross-link ECM components, and mediate cell-ECM adhesion (38, 70, 86). SPON-1 is spondin, a component of the ECM (44). VAB-19 is an ankyrin repeat protein, homologous to the human tumor suppressor Kank (76). VAB-19 is a cytoplasmic protein that localizes with components of epidermal attachment structures. Loss of any of these three genes causes defects in attachment of muscle to epidermis (29, 76, 90) and also results in increased regeneration (15, 29). These data suggest that muscle-epidermis attachment via the basement membrane inhibits regeneration. ECM proteins could act as a physical barrier to regeneration or could sequester essential soluble factors, rendering them unavailable to regrowing neurons. Alternatively, there may be receptor-mediated inhibitory mechanisms in C. elegans that are similar to inhibition of regeneration in vertebrates by CSPGs.
C. elegans neurons are capable of reconnecting directly to the severed distal fragment (14, 28, 57). Fusion restores membrane and cytoplasmic continuity between the two halves of the axon (28, 57). Fusion can also prevent degeneration of the distal fragment but only if it occurs relatively quickly after axotomy (57). If regenerating axons are given a choice of two distal fragments, one from the regenerating axon and one from another neuron, fusion occurs nearly exclusively with the correct fragment (57). Thus, reconnection after fusion can restore neuronal circuitry, at least in terms of morphology.
Fusion after axotomy requires the fusogen eff-1 (28). eff-1 also functions in homotypic fusions during normal development of epithelia, muscles, and sensory dendrites (54, 62). Fusion is also promoted by calcium and cAMP via PKA (28). Thus, fusion after axotomy is mediated by some of the same mechanisms as axon regeneration.
What of functional recovery? Regenerating axons in vertebrates can form de novo synapses and restore function providing they reach their target (1, 80). In C. elegans, loss of GABA neuron function (for example, in mutants that cannot synthesize GABA) leads to a characteristic shrinker behavioral phenotype (51). Severing all GABA commissures also results in the shrinker phenotype, and regeneration can restore GABA-mediated behavior (21, 96). Further, reduction of GABA regeneration (by increasing Notch signaling) also reduces behavioral recovery (21). Thus, regeneration can restore function to damaged neural circuits.
In a relatively short time, C. elegans has emerged as a highly tractable model to study axon regeneration. Current data suggest that many genes function similarly in worm and mammalian regenerating axons (see Introduction above). C. elegans offers the axon regeneration field a simple, highly invariant nervous system, genetic tractability, and the ability to study individual neurons in vivo. Although many novel signaling pathways that function in regeneration have been identified, many questions remain.
MAPK pathways (48, 95) and probably also Notch signaling (21) are activated by injury. The cellular mechanism that links injury to activation is currently unknown. A variety of injury signaling mechanisms have been proposed, including calcium entry, electrical signals, and changes in trafficking of a regeneration factor. Of these, only calcium is known to function in C. elegans regeneration (28). One possibility is that calcium signaling is the single cue that activates all of the acute responses to injury. For example, modulation of calcium in vivo and at physiological conditions can activate Notch (69). Alternatively, other injury signals may exist. For example, disruption of the microtubule cytoskeleton in nerve injury may be an injury signal. In support of this idea, microtubule depolymerization can activate dlk-1 signaling (9). However, it remains to be determined whether this activation mechanism occurs in regenerating axons.
Answers to these questions and others may come via genetic screens. So far, two screens (an unbiased RNAi screen in unc-70/β-spectrin mutants and systematic screening of existing mutant alleles using laser axotomy) identified many genes involved in regeneration (15, 36). Additional genetic screens are likely to provide more details on the function of known pathways and also to identify additional factors that mediate regeneration.
In the end, regeneration signaling must converge on growth mechanisms that enable the injured neuron to generate a new growth cone, maneuver the growth cone to its target, and reconnect. These growth mechanisms are for the most part not understood in C. elegans. However, the recent description of microtubule dynamics in regenerating axons and the discovery of a novel regulator of these dynamics (see section on efa-6 and Microtubule Dynamics) suggest that it will be possible to analyze regeneration at the cell-biological level. Further application of cell-biological techniques, such as electron microscopy and super-resolution imaging, in combination with genetic analysis, should result in a better understanding of the growth mechanisms that mediate regeneration.
A second approach to cell biology that also has potential translational applications is the use of C. elegans as a platform to screen for drugs that are effective in improving regeneration after injury. A high-throughput screen using microfluidics identified a chemical enhancer of regeneration (75). Further, microinjecting compounds directly into the pseudocoelom of animals immediately after axotomy can enhance regeneration (15, 21). This technique may be used in the future to assess limited numbers of compounds for effects on regeneration. Such compounds may identify particular cell-biological processes that are important for regeneration.
Most regeneration studies in C. elegans have used the morphology of the regenerating neuron as a measure of regenerative success. To date, two studies in the GABA motor neurons have shown that regeneration is accompanied by functional recovery at the level of whole-animal behavior (see section on Fusion and Functional Regeneration). However, such experiments are limited in their ability to accurately assess the function of individual regenerated neurons compared with their uninjured counterparts. C. elegans is a tractable model for a more detailed study of neuronal function using electrophysiological and optogenetic techniques (23, 33, 72). The future application of these techniques to the study of regeneration will yield information about how effective functional recovery is and may identify new pathways that are required for functional, rather than merely morphological, regeneration.
Axon regeneration can restore function, so why inhibit it? In the vertebrate CNS, pathways that inhibit regeneration also function in uninjured nervous systems and help maintain a stable, functional system by inhibiting aberrant growth and plasticity (2, 97). For example, CSPGs inhibit regeneration after injury in the CNS (11, 24, 50, 79, 97). In uninjured animals, enzymatic degradation of CSPGs results in ectopic growth and sprouting (5, 12, 20). These data suggest that inhibition of the injury response in the CNS is part of a broader program to limit plasticity. Similarly, it is possible that the C. elegans pathways that inhibit regeneration also function to stabilize the mature, uninjured nervous system.
Neurons in old C. elegans animals show a loss of stability and accumulate ectopic branches (85). Loss of jnk-1 causes an increase in ectopic neuronal branching in old animals, suggesting that jnk-1 contributes to nervous system stability (85). Consistent with the idea that stability pathways can also inhibit regeneration, jnk-1 mutants also display increased regeneration after nerve injury (36). In other ways, however, stability and regeneration seem to be separate processes. The dlk-1 pathway is required for regeneration but not for spontaneous branching in old animals. Loss of mlk-1 increases the incidence of age-dependent branching but decreases regeneration (36, 60, 85, 95). This suggests that in C. elegans, regeneration and spontaneous branching are promoted by different mechanisms. It remains to be seen whether other inhibitors of regeneration, such as Notch (21) and efa-6 (15), affect spontaneous branching and the overall stability of the nervous system.
Neurons in C. elegans can regenerate in response to injury. Is this ability due to a specialized regeneration mechanism that has evolved specifically to respond to nerve injury? Or is the response to injury part of a more general mechanism, such as homeostasis or stress response? Or is it even a pathological and undirected response to trauma?
During their life span in the wild, worms are subjected to desiccation, mechanical trauma, and predators. Thus, the ability to regenerate neurons quickly after they have been severed and to regain movement may pose a significant selective advantage. Alternatively, neuronal regeneration may be a particular manifestation of some broader biological process. A more complete understanding of the mechanisms that mediate regeneration might eventually help answer this question.
C. elegans has advanced the field of axon regeneration by providing a genetic system that is compatible with both high-throughput screening and single-neuron analysis. Additional forward genetics, a wider use of high-throughput techniques, functional regeneration assays, and drug validation will make C. elegans an even more robust model and further advance the study of axon regeneration.
The authors are not aware of any affiliations, memberships, funding, or financial holdings that might be perceived as affecting the objectivity of this review.
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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Happy (Dead) Presidents Day!
On this President's day, let's take some time not to look back at Lincoln, Jefferson, or that lovable walrus, Taft. Instead let's give some presidential props to socially conscious hip-hoppers Dead Prez, who asked the question:
You would rather have a Lexus? or justice? a dream? or some substance?
A Beamer? a necklace? or freedom?
(Lyrics and Word Cloud after the jump.)
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- Enrique Iglesias, Pitbull
- Micky Dolenz, Felix Cavaliere
- Mirage, Dog N Butterfly
Uh, uh, uh, 1-2, 1-2
Uh, uh, 1-2, 1-2, uh, uh
All my dogs...
It's bigger than..hip..hop..hip..hop..hip..hop..hip..
It's bigger than..hip..hop..hip..hop..hip..hop..hip-hop
Uh, one thing 'bout music when it hit you feel no pain
White folks say it controls yo' brain
I know better than that, that's game
And we ready for that - two soldiers head of the pack
Matter of fact, who got the gat?
And where my army at? Rather attack and not react
Back to beats, it don't reflect on how many records get sold
On sex, drugs and rock 'n' roll
Whether your project's put on hold
In the real world; these just people with ideas
They just like me and you when the smoke and camera disappear
Against the real world *echos*
It's bigger than all these fake-ass records
When po' folks got the millions and my sista's disrespected
If you check 1-2, my word of advice to you is just relax
Just do what you got to do; if that don't work, then kick the facts
If you a fighter, rider, biter, flame-ignitor, crowd-exciter
Or you wanna jus' get high, then just say it
But then if you a liar-liar, pants on fire, wolf-crier, agent wit' a wire
I'm gon' know it when I play it
Uh, who shot Biggie Smalls?
If we don't get them, they gon' get us all
I'm down for runnin' up on them crackers in they city hall
We ride for y'all - all my dogs stay real
Nigga, don't think these record deals gon' feed your seeds
And pay your bills, because they not
MCs get a little bit of love and think they hot
Talkin' 'bout how much money they got; all y'all records sound the same
I'm sick of that fake thug, R&B-rap scenario, all day on the radio
Same scenes in the video, monotonous material
Y'all don't here me though
These record labels slang our tapes like dope
You can be next in line and signed; and still be writing rhymes and broke
You would rather have a Lexus? or justice? a dream? or some substance?
A Beamer? a necklace? or freedom?
Still a nigga like me don't playa-hate, I just stay awake
This real hip-hop; and it don't stop 'til we get the crackers off the block
They call it...
[Hook 2x]
[Repeat 6x]
D.P.'s got that crazy shit
We keep it crunked-up, John Blazed and shit
(*"They call it, call it, call it" -> stic.man*)
(*"Fake, fake, fake records" -> M1*)
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Capital University film studies
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Information about Capital University film studies, and registering for discussion sections and departmental electives. As a student, your first stop should be the registrar's office, where you can pick up a free copy of both the academic calendar and course catalog, in order to find available credit classes that you may be interested in taking. You may be surprised that you can enroll for a full-time course load, without having to attend any 8am classes at all. Finally, check with your major department for assistance with prerequisites, tuition payment deadlines, and Capital University Film Making information. Please use the form above to request admissions info for Capital University.
Becoming a professional film actor is a very difficult process. Improvisation allows movie actors true expressive freedom, since they themseleves don't know how the situation is going to turn out. In professional acting, improv is also used to cover up momentary lapses, so the audience isn't startled. The actor?s performance is mediated by particular performance signs including facial expression, emotion, and vocabulary.
Classical acting is an umbrella term for a philosophy of acting that integrates the expression of the body, voice, imagination, improvisation and script analysis. Method acting by comparison, is a range of techniques based on for training movie actors to achieve better characterizations of the characters they play. Part of the training includes actors memorizing lines and therefore being able to work without a movie script. Other acting classes may include mask work, improvisation, and showing emotion for the camera. Beyond consideration for Capital University film studies, you may think about Sinclair Community College major list for complementary electives to your primary course of study.
Admissions Counselor I
University of Texas at El Paso
Composes correspondence to answer admission inquiries. Maintains and enters admission information into database.
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Works collaboratively with Admissions for re-enrollment. Maintains tracking record of seniors college choices..
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Experience working with college admissions. Management of admissions criteria, policies, and student assessment.
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The person should be an energetic and enthusiastic individual who will articulate admissions policies and procedures to students, parents, and counselors.
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College of DuPage
Coordinate outreach efforts with Admissions and Marketing to identify and recruit students into the S-Stem program.
7 days ago


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27 Pages Posted: 1 Nov 2011
There are 3 versions of this paper
Date Written: August 1, 2011
This paper models policy responses to changes in solvency by Dutch occupational pension funds using a unique panel dataset containing the balance sheets of all registered pension funds in the Netherlands over a period of 15 years (1993-2007). The model describes how nominal pension rights are expanded, by e.g. indexation or backservice, or, on the contrary, how the current pension accumulation is skimmed, e.g. by setting the pension premium over its actuarially fair price to build buffers. Policy responses are explained by the funding ratio and other pension fund characteristics such as pension funds' size and type, and participants' ages. We find that pension rights are expanded in line with the funding ratio, but that the pension funds' response function exhibits two sharp and significant behavioural breaks, close to the minimum funding ratio of 105% and the target ratio of around 125%. These levels also play a pivotal role in current supervisory regulation. We further find that large pension funds and grey funds are relatively generous to participants.
Keywords: pension funds, pension rights, risk sharing instruments, indexation, funding ratio, solvability, regime shifts
JEL Classification: G23, G28
Suggested Citation: Suggested Citation
Bikker, Jacob Antoon and Knaap, Thijs and Romp, Ward E., Real Pension Rights as a Control Mechanism for Pension Fund Solvency (August 1, 2011). De Nederlandsche Bank Working Paper No. 311. Available at SSRN: https://ssrn.com/abstract=1951909 or http://dx.doi.org/10.2139/ssrn.1951909


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As part of my course, we have to put on an exhibition. As a class we are putting on a 3 part exhibition revolving around the theme of 'Place'.
The opening night is the 24th of March, at Out Of The Blue Drill Hall, but parts 2 and 3 open on the 31st of March and the 8th of April respectively, with a big and boozy closing event on the 11th of April.


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Day 14, social activitiesAug. 14th, 2010 05:39 pm
Since I'm about to head out of the house for 20-40 hours of parties and other sundry social events, I figured that sort of thing would make a good post. I am involved in a lot of social circles, covering a huge cross-section of different subcultures in and around Atlanta, so if you're bored then read on and find something new to explore. I just hope I don't insult anyone along the way...
( Read more... )
( Read more... )


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Sun, Jul 23 Frankie's
Time 8:00 pm
Time 8:00 pm
The Louisiana Joymakers, a Dixieland jazz band, started life in 2002. Since then they have played in numerous festivals, clubs and other venues in Canada and the US. Jim Armstrong fronts the band on trumpet and vocals with Ben Henriques on reeds, Clive Collins on trombone, Steve Toren on tuba, Scott Robinson on drums and Mike Hobbs on banjo and guitar. Their high energy music, a consistent hit with swing dancers, is mainly from the 1920's and 30's with some more modern tunes thrown into the mix.


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Responsible Trap-Neuter-Return (TNR) IS a vital part of keeping community wildlife safe. Without responsible TNR cats would continue to reproduce and be forced to hunt for sustenance. Remember, members of your neighborhood that engage in TNR care about wildlife too and took it upon themselves to help implement an effective solution to reduce and control outdoor cat populations. Feral cat feeding stations encourage outdoor cats to eat cat food, and a cat whose stomach is full is less likely to hunt wildlife.
The following are suggestions for keeping wildlife safe and for keeping cats away from your property if that is a concern. None of these methods will harm animals. If you are interested in deterrents for your garden or other areas, see:
http://www.alleycat.org/deterrents or http://bestfriends.org/resources/solutions-cat-related-issues
Bird Feeder and Bird Bath Placement
- Keep feeders and baths at least five feet (ideally 10-12 feet) from shrubs, trees or cover that can conceal a stalking predator.
- Avoid using low feeders or ground feeders that make it easier for predators to capture wild birds.
- Use plastic or metal poles to support feeders so predators' claws cannot help them climb to the feeder.
- Fix a downward opening cone or cookie tin to the bird table post to prevent predators climbing up. Squirrel baffles are a good choice. Spiked tree collars can also prevent climbing up a tree. Other tree collars can be made from slick plastic or metal.
- Spiny plants or clippings, such as thorned mesquite, cactus, ocotillo, or an uncomfortable surface around the base of feeding stations to prevent predators from sitting underneath.
- Clean up spilled seed regularly to minimize ground feeding birds.
Bird Houses & Nesting Boxes
- Choose designs with steep roofs and lacking perches to help deter predators.
- Keep nesting boxes at least eight feet off the ground to decrease predators jumping to the box.
Property Perimeter and Fences
- Repair or block gaps in fencing that may be access points for predators.
- Use flimsy plastic barrier fencing mesh (NOT netting! Lizards can get caught and die in nets) placed on top of a fence (usually between poles made from something such as PVC or garden stakes) to prevent cats climbing over it. It is available in green and brown. An outward-angled chicken-wire fence will do the same thing but PLEASE check daily for lizards.
- Specially designed strips of plastic spikes, such as Cat Repeller Fence and Wall Spikes, can be placed on top of a fence, shed roof and nest-box roof preventing predators from walking on them. We discourage anything metal because lizards, birds and other wildlife may be harmed.
Trees, Shrubs, Hedges, Brush, etc.
- Decorative or wire fencing installed in front of shrubs or hedges that may conceal predators can prevent attacks from these vantage points.
- Plant wildlife-friendly vegetation, such as prickly bushes to provide secure cover for birds. These should be close enough to where birds feed to provide cover, but not so close that predators can use it to stalk birds. This kind of foliage may also provide food and nesting sites.
- Brush piles and shrubbery should be checked regularly for ground nests and fledgling birds that are most vulnerable to prowling predators.
Make a "Safe" Area
Surround a smaller area with a fence, such as chicken wire, that leans in the direction of the predator approach which will make it impossible to climb over due to the angle.
Ultrasonic Deterrents
The effectiveness of these is probably variable, and there is little scientific evidence of their effectiveness. Some are also quite expensive. It's a matter of individual choice whether to use them or not.
Aim a motion-detecting ultrasonic device at the area where predators enter your yard. It emits a high-frequency sound that humans can't hear. The loud sound is unpleasant to the predator.
- Essential Oils
- Citronella: Mix 1 part Citronella oil to 4 parts water and spray liberally in the trouble spot.
- Lavender: Mix 1 part Lavender oil to 3 parts water in a spray bottle and shake well. Spray liberally in the trouble spot.
- Lemon or Orange: Scattering lemon or orange peels on the ground in the off-limits areas. Squeeze a lemon into a liter of water in a squirt bottle, add 2 dozen drops of eucalyptus oil and shake well.
- Peppermint: Mix 1 part peppermint oil to 3 parts water in a spray bottle and shake well. Spray liberally in the trouble spot.
- Mix 2 tablespoons of Cinnamon, Rosemary and Lavender into a pint of boiling water and leave it overnight. Next morning you will need to strain the liquid through a cheesecloth and then add half a cup of vinegar and a dozen drops of Tangerine essential oil. Place the liquid in a spray bottle and shake well. Spray liberally in problem areas.
- You can also try soaking cotton balls in one of the oils and putting them in and around the area.
- Rosemary is a perennial that you can plant which grows well here, smells nice, is useful for cooking and cats hate the smell.
- Coleus canina can be planted about three feet apart around the bird feeder emits an odor cats find offensive, making them think twice about returning to your yard. I don't know anyone that has been able to grow this in Tucson however. It is marketed under the names Pee-off and Scaredy-cat.
- Use a product called Silent Roar to scent mark your garden
Install a motion-detecting sprinkler system. Aim the motion detector away from the bird feeder to the area where predators enter your yard. When the system detects a cat, it activates and squirts water in that direction, which scares the cat away.


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gnat(redirected from GNATS)
Also found in: Dictionary, Thesaurus, Medical, Acronyms, Encyclopedia, Wikipedia.
Related to GNATS: fungus gnats
down to a gnat 's eyebrow
Fig. down to the smallest detail. He described what the thief was wearing down to a gnat's eyebrow. No use trying to sneak anything out of the refrigerator. Mom knows what's in there, down to a gnat's eyebrow.
strain at gnats and swallow camels
Prov. to criticize other people for minor offenses while ignoring major offenses. (Biblical.) Jill: Look at that. Edward is combing his hair at his desk. How unprofessional. Jane: Don't strain at gnats and swallow camels. There are worse problems than that around here.


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This restaurant is a gem , it offers beautiful dishes that are done with passion . It is my second time and i just love the place . It responsibly priced .
1 - 10 of 22 reviews
This restaurant is a gem , it offers beautiful dishes that are done with passion . It is my second time and i just love the place . It responsibly priced .
Gambas croustillants were great for starters, and had the thon mi cuit as a main. Both were very good and the main was copious. Worth another visit
We were staying on Rue d'Aboukir and looking for somewhere for a wedding anniversary meal - with the usual Paris problem of being vegetarian. La Mauvaise Reputation was the first place that we looked at the menu at and saw that they had a couple...More
I won't spend too much time going over the same territory that the reviews have already covered that led us to come here ourselves, but suffice to say that we also found the food to be incredibly delicious, the staff to be charming and welcoming...More
I have been here already several times as it is close to our Paris office. Food is always good and fresh. This time we took Salmon Maki as a starter and than Risotto with mushrooms and safran. Really tasty. Make sure you take Coffee gourmet....More
Sometimes you just feel a place will be great. Rarely, it truly is. The Mauvais Reputacion is great. Wonderful flavors all made in house and service that was mellow and attentive at the same time. A win.
I went there for a dinner with my colleague. Service was very professional and fast. We relayed on waiter recommendation and we didn't made a mistake. Food was really good which I can also say for the red wine. Price is ok for the Paris...More
Great place,informal,excellent food,good size not a spot on the dish.Good wine,will be back next time in paris
To be frank, I can't add much to what has already been said below. The previous 10+ reviews are – without exception – accurate and positive. On two visits to Paris we've dined three times at La Mauvaise Reputation – this is a restaurant that...More
Went to this resto with friends during the week. Excellent service including an informal visit from what looked to be the chef. Set menu with interesting combinations that were extremely well presented including pork from Bigorre with cepes and potatoes and ended up with a...More


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The Azulita Project was born from the desire of several individuals to create a healthier and more environmentally sustainable community and world. Moved by the rampant plastic pollution and overuse of chemical pesticides in the Los Llanos region, the four original founders—Mike, Lainie, Erwing, and Pato—joined forces in 2008 to create the Azulita Project.
In its initial form, the Azulita Project was a volunteer "communitarian" effort which included organic farming education programs, a recycling project, a youth soccer league, dance instruction, English classes and a community space for teaching and gathering.
In 2014 the Project founders reorganized the recycling portion of the project in pursuit of an effective and sustainable business model. The concept was simple: create an economic incentive for people to recycle plastic.
Azulita team members purchase plastic waste from local residents, clean the materials, and create bales of crushed plastic containers. When the Azulita plant has compacted between 10 to 15 tons of waste plastic—equivalent to a semi truck load—the bundles are delivered to the city of Guadalajara where they are sold to a recycling facility. The proceeds from plastic sales are reinvested into the business, the community, and other Azulita initiatives.
The Azulita recycling program actively removes up to 1 ton of plastic waste a day from the Los Llanos region. Prior to the Azulita Project, this waste would have been burned or tossed away, polluting local beaches and, ultimately, the ocean.
Others have attempted to start recycling programs in the region but have failed for various reasons, including lack of local interest and sustained motivation by residents and visitors.
The Azulita Project recycling program is successful due to its innovative, incentive-based model that has dramatically increased local recycling participation. Inspired by the effectiveness of deposits on glass bottles and the value of various metals, Azulita created a privately subsidized buy-back program that places value on post-consumer plastic and encourages residents to ollect waste plastic and deliver it to the Azulita facility.


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Ohio Hardwood Furniture
Photo 1 of 7Next
It and various aged table seats might also combine. Things for example platforms yard / huge potted flowers terrace, and seats also can enhance the sweetness of the inner of the old house that is house.The isn't such as a home nowadays. The team of area sometimes appears unusual. Whilst the bedroom is very slim, eg so spacious family room.
Thus will be the home which will be extended. Properly, you are able to work this around by the addition of a Ohio Hardwood Furniture in a space that's too extensive or switching characteristics. For instance a lot of the kitchen along with room, while half the room used as being a storage


text:
Call us @ +1-888-483-3317 for Windows 10 Technical...
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Everyone @ Suzanne Brown has always been wonderful & professional....recently Meredith ********* was a super star who got my new vehicle added onto my policy instantaneously and answered all my questions....soooo blessed to have her!
Wow, Suzanne. Thanks for your efficiency and expertise. Very happy with our decision.
Thanks so much!
Awesome!!! You do good. T.J. was bragging on you last night with your service concerning his accident details. You are our super-insurance hero!!
We had a very pleasant experience, honestly better than any other in the past.
We would recommend this agency and will use them again next year for renewals.
Of those that I contacted, the Suzanne Brown agency not only took time to determine what was best for me, but spent time to fully inform me of my options. I realize that agents do have more than me as a customer, but the agent made me feel as if I was of great importance to her and the agency she is with. As I tell my team, it only takes a few minutes to make a customer or prospective customer feel important. I expect that of my team and I definitely expect that of someone I do business with. If you are in the need for insurance, I highly recommend that you ask for this agency. They are very professional and timely in response, making sure that your needs are adequately met.
Spoke to the claim officer at ASI, and they said they would be dispatching a company that specializing in smoke/fire damage clean up, a cleaning service company that tries to clean sofas, pillows, etc., and also an electrical engineer to determine the root cause of the fire.
A loss adjuster was also able to get out there on site today.
So far I am blown away by the responsiveness of ASI.


text:
Copiah-Lincoln Community College admission interview
Think about what career is best suited for you, and where you want to study. Request more information now.
Think about what career is best suited for you, and where you want to study. Request more information now.
Think about what career is best suited for you, and where you want to study. Request more information now.
Find info concerning Copiah-Lincoln Community College admission interview, and registering for discussion sections and department seminars. Which college degrees give you the best chances of finding employment after graduation? Concentrating on Copiah-Lincoln Community College admission interview, for example, may give you an advantage in corporate recruiting being conducted on-campus. Check with the admissions office for course registration deadlines, and college admissions program requirements. Please use the form above to request admissions info for Copiah-Lincoln Community College.
Before an admissions interview, you should prepare answers for difficult interview questions that may be asked. Knowing what to expect when going for your interview can set you apart from other applicants. Interview questions are generally grouped into several broad areas, such as motivation and personal development. First impressions can set the tone for the rest of the interview, so focus on building rapport in the first few minutes. Finally, you may be asked technical questions that relate to concepts that you studied in your major, so organize your thoughts and show an understanding of the issues.
Write out answers to questions you think the admissions committee will ask, as well as a list of your strengths, weaknesses, and academic accomplishments. Your main goal in an interview is providing the interviewer with detailed past examples of your achievements. Further, you need to practice your acting skills, so look good, speak clearly, and act like a confident, professional person. Finally, be honest, and consistent in your answers, and remember that you are being evaluated at all times. In school activities and during meals, your ability to interact with people may be observed. Beyond major requirements within the admissions department, you may consider Marygrove College preparation for complementary electives to your primary course of study.
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Centura College
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Salon Institute
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Portland State University
The Graduate Admissions Evaluator will support Graduate Admissions in the Office of Graduate Studies through coordinating with departments and processing..
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UC San Diego
Knowledge of UC policies and procedures related to graduate education including admissions, diversity, student services, and other related topics.
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Yeshiva University
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Hello, I am trying to Sell my 01 YZ426F, so I tried getting it started today which is where I ran into a problem. The bike would only start with the choke on and would not idle at all... I had to have the throttle open for it to run. I tried cleaning the air filter, I replaced the spark plug, and put brand new premium gas in it. Still it was doing the same thing. When it was running it was sputtering and backfiring like crazy! Anybody have any sugestions for me?!?!
I need help!!!! Yz426f
Sitting a while, eh? Clean out the pilot jet.
What he said!!!!
just take the jets out and blow some compressed air thru 'em. refit and check that.
check the pilots
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I didn't have high expectations for this place but I was still extremely disappointed upon arrival. We had read online that most rooms were lake view rooms. This is false. Most are NOT. We had a ground level room & there was a woman that walked past our room a few times & would stop & glare in on us through the window, which was totally creepy. We had to keep the curtains drawn in an already dreary atmospere. The room smelled terrible & I could barely breathe. I had to wash all of my clothes twice when I got home to get the stench out of them. As we were getting ready to get out of there ASAP the next morning...We noticed a pile of either dog or cat food under the desk. We had brought neither. Which could only mean that the housekeeping definitley wasn't up to par. I don't even want to think about what was on the bedding. Also, the a/c unit was so loud that neither of us slept. It was a HUGE disappointment. Do not stay here! Cheap & there's a reason why!
- Official Description (provided by the hotel):
- Located near the shores of Lake Superior in Duluth's historic Old Downtown. Walking distance to shopping, restaurants, theaters and entertainment, Canal Park, Fitger's Brewery Complex, Fon du Luth Casino, as well as major hospitals and clinics. Full Cable and Showtime, bring your laptop - we have wireless internet access! Standard rooms, some with lake view, whirlpool lodge rooms with fireplaces, rough log furniture, pine walls and wonderful views of Lake Superior. Free continental breakfast and a friendly staff. Enjoy your stay at Voyageur Duluth Lakewalk Inn! ... more less
- Reservation Options:
- TripAdvisor is proud to partner with Travelocity, Booking.com, Priceline, Hotels.com, Expedia, Orbitz and Agoda so you can book your Voyageur Lakewalk Inn reservations with confidence. We help millions of travelers each month to find the perfect hotel for both vacation and business trips, always with the best discounts and special offers.
- Also Known As:
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text: Seller: sprats14 (174) 100%, Location: Hereford, Ships to: GB, Item: 322331701664 Condition: In good condition with usual marks commensurate with age and use., Product Type: Display Cabinet, Original/Reproduction: Original, Style: Edwardian, Age: 1900-1950, Material: Mahogany, Details: LOVELY EDWARDIAN CORNER DISPLAYCABINET DIMENSIONS: Height: 52.75" / 1340mm Width: 24.25" / 616mm Depth: 16.5" / 419mm CONDITION: Very clean, Sturdy Cabinet, all Joints Good, Hinges Good, Strong Legs, Possible Lower shelf replacement, Original Glass, Lock with Key. *PLEASE CLICK IMAGES FOR LARGER PICTURES* Please Email or Telephone Andy with any questions on : 07979 650044 DESCRIPTION: This Attractive Edwardian Corner Display Cabinet is in Great conditon having an Inlaid Galleried back, above a Crossbanded Frieze and a Single Bow Fronted Inlaid Glazed Door with Original Glass, fitted with a Lock to which there is a Key, opening to reveal a fitted Interior with One Removable Shelf, the Inlaid Stiles either side of the Door are adorned with Flower Design, all standing on Square Tapering String Inlaid Outswept Legs united by an Undertier, which possibly is a Previous Professional replacement. A great addition for the Home to display those Prized possessions. * SEE ADDITIONAL IMAGES BELOW* DELIVERY INFORMATION: Delivery for this item via courier is £45.00 to Mainland England and Wales, but please contact us in the first instance. Please contact us for prices for delivery to Scotland and Europe. *Terms and conditions apply* CONTACT INFORMATION: Please contact Andy on 07979 650044 for any information required or alternatively email us at firstname.lastname@example.com. Thankyou for viewing - Please bid with confidence. ADDITIONAL IMAGES:


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Three men charged with terrorism offences have been remanded in custody.
Yousaf Shah Syed, 19, from High Wycombe, Bucks, Nadir Ali Sayed, 21, from Hounslow, and Haseeb Hamayoon, 27, from Hayes, today appeared at Westminster Magistrates' Court charged with the intention of committing and preparing for acts of terrorism.
The trio were arrested in a series of overnight raids in outer London and the Thames Valley area on November 6 and 7.
The men were remanded in custody until their next court appearance at the OId Bailey on December 4.
No application for bail was made.
Properties in Hounslow, High Wycombe, Uxbridge, Southall, Greenford and Hayes were searched as part of the raids.
A fourth man, arrested in Uxbridge, was released on November 15 without charge.
No further action is being taken against him.


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Five more people have died in Hennepin County from carfentanil-related overdoses, according to a May 19 report from the Hennepin County medical examiner.
Eden Prairie resident Adam Patrick Hall, 32, was one of the five people who died. Hall died April 17, early in the morning at his residence on the 6000 block of Kingfisher Lane, according to the release. The report notes the cause of death as mixed drug toxicity (carfentanil, fentanyl, methamphetamine).
"We remain vigilant and continue to order the carfentanil and designer opioid testing on a case-by-case basis after considering the autopsy findings, investigation and laboratory results," Dr. Andrew Baker, Hennepin County medical examiner, said.
Officials have expressed concern over the increasing prevalence of Carfentanil, a powerful new opioid to the state. This spate of five deaths brings the total of deaths related to the drug countywide to 10.
Some 10,000 times more potent than morphine and 100 times more potent than fentanyl, the drug is intended to be used as a tranquilizer for elephants and other large animals. Carfentanil requires specialized laboratory tests to detect, slipping past routine drug and alcohol screens, and can even be harmful to first responders if they are exposed to it.
The other victims noted in the medical examiner's report were a 26-year-old Minneapolis woman, a 39-year-old Minneapolis man, a 23-year-old Maple Grove man and a 43-year-old Plymouth man.


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Wet March breaks Broken Hill average
Bureau of Meteorology statistics have revealed the March rainfall in Broken Hill has topped 90 millimetres more than average.
Broken Hill recorded 112.4mm of rain for the month of March, with the highest daily rainfall of 96.6mm falling on March 5.
The annual rainfall total to the end of March is 225mm.
The mean maximum temperature in the city during March was just below average, with a temperature of 28.1 degrees. However, minimum temperatures were slightly above average at 16.2 degrees.
The highest temperature recorded in Broken Hill for the month was 35 degrees on March 27, while the lowest temperature was 9.4 degrees on March 10.
Topics: rainfall, weather, broken-hill-2880


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This is a super easy and quick tutorial for these fun red and gold polka-dot Mickey nails, perfect for the holidays or simply for the fall/winter! I went to work with them for one day and already got a compliment, so I'm hoping you will too! All the supplies are very inexpensive and not hard to find either. You don't need any special training for this look, just some cheap household items and 15 minutes or so! Please keep in mind that I am not a professional or anywhere near a hand model so please bear with me!
Gather these supplies to start:
- Base coat nail polish
- Essie nail polish in'Life of The Party'
- Gold glitter nail polish (does not have to be a nail art brush)
- Regular clear top coat nail Polish
- Bobby pin
- (Optional) acetone sponge jar and/or cheap dollar store paint brush
Now that we have all our supplies, it's time to get down to business… (to defeat the Huns).
We'll start with clean nails, topping them with your regular base coat of polish.
One coat down of the Essie nail polish in the shade 'Life of The Party'. I believe this shade was part of a holiday edition to boot! If you don't want to buy new nail polish that's totally fine, just use a darker red that you have in place of this shade!
Coat 2 and we're almost there! I realise that the polish got on my skin in some places, bare with me and I'll show you a secret!
For the next step you'll need the bobby pin and the optional cheap paint brush. You're going to take the bobby pin and dip the little ends (like in the picture above) in the gold glitter polish.
Then, in the center of your nail, make 3 small dots with the end of the bobby pin using the gold glitter polish to make a little Mickey head.
Continue with the bobby pin (or don't if you like the simple look), going along with the shape of the Mickey head to make your other dots like the picture above. Try not to put them too close in order to keep them from turning into a giant glittery blob!
Now after you finish the dots, you may want to clean up around the nails. This is where that cheap brush comes in. Try to find a very small (preferably hard bristled) paint brush at the dollar store or other cheap outlet. Cut the end of the bristles at an angle and dip the brush in the acetone sponge for a quick and inexpensive nail polish removing brush! I can't tell you how much quicker and easier it is than a cotton swab or waiting for the nail art barrier to dry!
And there you have it! Let them sit for a minute or so, finish it with a top coat. Easy-peasy right? Now your nails will be festive and fun with out the hassle and cost of getting them done!
Let us know how you liked this tutorial and if you tried it! I'd love to hear about your versions!


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CO Barista Freaks Out Over Unicorn Frapps
I so feel this dude's pain. I was a short-order cook for a long time before I got into radio, and if you've ever worked in the restaurant (or related) industry, you know: Sometimes, all reason and logic cease to exist, rendering you a sniveling, resentful, self-loathing slave who wonders why you were ever born. Other times, however, the work is extremely enjoyable.
Behold, one of Colorado's own, giving his thoughts on Starbucks' new Unicorn Frappuccino:
Subscribe to 94.3 the X on
The news got in touch with him, and he said he didn't mean to criticize the drink. And he's not in trouble with Starbucks . . . they said they'd reach out to him to see if they can improve his experience.
Respect, bro.


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Thanks to Tom for transcribing these.
During the summer of 1965, Paul and Mark spent time in France living with two different families. Upon their return, their teenage-boy counterparts from these families ventured to Brown Street for a while. (Maybe someday Paul M will provide more details and reminisces). Below are Mark's letters home during his stay. He stayed in Le Vesinet, an upscale suburb of Paris, whose subway stop was the next one beyond where Dick got off each working morning during his Paris assignment, nearly 35 years later.)


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Before I was a Kidney Warrior I was a Marine
End-Stage Renal Disease (the last stage of kidney disease) is a medical condition in which a person's kidneys cease functioning on a permanent basis leading to the need for a regular course of long-term dialysis or a kidney transplant to maintain life (source: cms.gov).
Kidney Disease is one of the lesser known illnesses. Maybe because kidneys are a pair of the lesser known organs in the human body. Personally, I knew the bare minimum about kidney disease because of a relative who received a kidney transplant after being on dialysis for a few years. I thought it was just an old person's/unhealthy overweight person's disease. It would never happen to me.
I felt the healthiest I ever did in my life! In 2011, I was in Afghanistan …I was a Combat Camera Marine. And then one day my legs and face began swelling. After a few weeks of being stubborn, I finally went to medical. In a matter of days, I was immediately medically evacuated to Landstuhl Regional Medical Center, Germany. Two weeks in Germany, then back to the states to Walter Reed National Military Medical Center. I got to spend a lovely amount of time there undergoing every test you could imagine to ultimately be diagnosed with an autoimmune disorder which caused my kidneys to fail.
This caused a bunch of "you can't make this stuff up" life experiences that changed my entire life. I received my second chance at life on April 2, 2011. My living donor is the most selfless, caring person that I have ever met…except that I didn't meet her until after she agreed to be my kidney donor. It took me feeling my life leave my body to reach out for help…and she found me.
When I woke up from the transplant surgery, when I realized that it was over I started crying. I was so happy. It could have been the drugs, but all I could think is that I wanted everyone who survives by dialysis to feel the way that I do right at this moment.
Would you like to share your story and be featured on Faces of Kidney Disease? Join the I Am The Face Campaign.


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Published on Jan 6, 2017
Up to 30 professional development contact hours (CEU/PDHs) available. TED has connected more than 500 teachers across the Atlantic since it was first launched in 2010!
Deadline for applications: January 15, 2017.
Register here soon!
There is no application or participation fee for TED. There is no cost to participate. TED is FREE!
Read teacher testimonials!
Are you interested in talking to other teachers about educational issues beyond classroom management? Do you get excited about topics such as immigration, youth culture, notions of the "other" in national identity formation, and education in rural vs. urban contexts? Are you curious about differences in education between the US and other developed (and developing) nations? If so, we invite you to join a 12-week online program: "Transatlantic Educators Dialogue, 2017".
This project is coordinated by the European Union Center at the University of Illinois at Urbana-Champaign. We are recruiting teachers in the US and Europe to come together once a week (Sundays 12noon-1:30 pm CST in the US) in online sessions to explore a variety of educational topics as they relate to issues of identity and difference. Participants will meet simultaneously online using the Elluminate application; in addition, prior to each session, participants are expected access an online course website to read materials and post responses related to the week's topic.
The "Transatlantic Educators Dialogue" (TED) program will begin on February 5, 2017 and will conclude on April 30, 2017. Online sessions are 90 minutes. Weekly attendance is not mandatory, but expected and helpful as discussion sessions will be more productive when a variety of perspectives are shared.
All participants enrolled in the program will receive a Certificate of Completion. Up to 30 non-credit professional development contact hours will be recognized:
Up to 30 non-credit Professional Development Hours (PDH) for Illinois teachers;
Up to 3.0 Non-credit University of Illinois Continuing Education Units (CEU) for 30 contact hours for non-Illinois teachers.
Please register online at TED website or by clicking here by January 15, 2017.
Please contact Lucinda Morgan (TED Coordinator) at firstname.lastname@example.com if you have questions about the program. Enrollment will be selective in order to ensure a diverse geographic participation.
For more information: http://europe.illinois.edu/ted/


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Speakers Bootcamp is for Public Speakers, Motivational Speakers, Coaches, Trainers, Consultants, Toastmasters and anyone who wants to follow the profession of speaking professionally.
Speak for High Fees and in Many Lands
Frank Furness is not only a marvellous, successful professional speaker, he knows all the insiders information about speaking globally. If you are dreaming and planning to become an international speaker, you will treasure and benefit from every word. I wish I had Franks great ideas before I made my first international presentations. It always pays to get expert advice from a successful expert who has been down the trail before you.
Dottie Walters CSP http://www.speakandgrowrich.com/
How Often Do You Find Yourself Saying: "I Wish I Knew How To Speak At Conferences And Conventions"
The Bootcamp will provide Proven Tips, Tools and Tactics To Speak At Conferences And Conventions.
Frank Furness, originally from South Africa has been living in London since 1993. He embarked on a full time speaking career in 1997. Since that time he has spoken in 48 Countries around the World and continues to secure International speaking assignments. Most of the engagements are self generated and Frank has become a master of networking and identifying opportunities. He runs his speaking career as a professional business. When Frank started his speaking career, he spent a lot of wasted time floundering around and trying to find out how to market himself.
Over the years he has developed professional relationships with some of the best marketers and speakers in the World and learnt from them. Two hours of personal coaching time with Dottie Walters cost Frank $500 and the advice he received resulted in his first tape course order for R600,000.
Over the last four and a half years he has established clients that are amongst the largest Blue Chip Companies in the world. Sponsorship and arranging his own speaking tours has contributed to his success and International exposure. Frank has now run three successful bootcamps and the delegates are raving. With the ideas learned on the bootcamp, one delegate secured 11 speaking assignments in the week following.
The idea of a BOOTCAMP originates in America where Frank has attended them and experienced the positive results. A BOOTCAMP is a weekend of intensive learning and action planning. The first day of BOOTCAMP will concentrate on the speaking and platform skills, while the second day concentrates on marketing, products and running a business
Discover how Frank has secured bookings in 48 countries - By following the proven branding and marketing strategies shared on the bootcamp, you too could be speaking globally or making a big name for yourself as a speaker in your country.
Develop and sell your own products - Frank shares his strategies that helped him to generate over 35 products and how his first ever product sale was for $70,000
Learn from the greats. - Watch videos, listen to interviews and discover the secrets of success of some of the worlds greatest speakers
I came to Frank's bootcamp as a successful speaker, TV & radio presenter and media journalist. I left with seven pages of notes & an understanding that I still had a great deal to learn. Wow!
Michael Jackson http://www.speakerpartnership.com/michaeljackson.htm
Learn how to run a speaking business - Find out how to run a successful business - branding, marketing, articles and securing bookings on an ongoing basis will be shared on day 2.
Develop a 'Killer' keynote -Frank shares the formula to developing a keynote that will have your audiences shouting for more.
Use multimedia to enhance your presentation -Find out how to avoid 'Death by PowerPoint and use multimedia effectively.
Learn the essential elements to any succesful presentation - Discover how to use the 7 'E's formula, humour, pauses and creating a magical experience for your audience.
Grow your business - Frank will show you a marketing kit that has a 95% success rate in securing bookings. He will show you how to build your profile and market yourself.
Get the bookings - Franks 17 strategies to get booked includes, sponsorships, bartering, famaliarity days,working with bureaus and much, much more.
Sam Storey - Richard Bevan - Gladstone Small - Dominic Cork
British Middleweight Boxing Champion - England Cricketers
Listen to the comments from the Sporting Bootcampers
Venues and Dates
10-11 March 2012
2-3 June 2012
19-20 May2012
11-12 August 2012
By attending the bootcamp, you will; Discover your uniqueness. Find your niche market. Learn from the worlds top speakers. Get constructive feedback from the other bootcampers. Develop a marketing pack and strategy. Learn how to produce and market products. Speak in exotic locations. Have fun and earn BIG BUCKS.
Take a look at some Bootcamp Photographs
United Kingdom * Singapore * South Africa * Malaysia * Olympians and Sporting Champions
The course was awesome. You demystified the mysticism surrounding professional speaking and yourself while enhancing your character, strength of personality and generosity of spirit. I have rarely seen anything like it in business. It was humbling and awe inspiring. Your techniques were from an incredible level of experience. You are a master speaker but also a master at getting work. A hundred books could not have done what you did in two days - show me how to make money from speaking. You also showed me how to have fun at the same time
David Thomas - Professional Speaker http://www.davidthomas.tv/
An intense experience which made me realise how much I still have to do to allow me to market myself well.
Marc Woods http://www.marcwoods.com/
I have no doubt that this has been the best two days in my trainer/speaker business. The course was simple, practical and effective. Thank you for taking the time to humbly share your experience & ideas that will certainly make a difference in my life and in my business. Frank you are an amazing man, one who is exemplary and worthy of emulation. Clearly you demonstrate a sincere and genuine interest to serve people & see them grow!
Paul Naidoo http://www.paulnaidoo.co.za/
After the bootcamp, if you follow all the strategies covered, you will be able to speak with confidence and have a fully booked diary of speaking engagements and live the life you love.
You will receive; 150 page workbook A resources CD with over $500 wort of information and software Lunch,and refreshmants both days Discounts on Frank's resources.
Money Back Guarantee
If you follow all the steps shared over the two days and can't make it as a speaker in the next six months, we will refund you and you can keep the Action planner and resources CD.
Book now as this bootcamp will only happen once a year in your country. You can't afford to wait...
Bootcamp Dates and venues
London, United Kingdom - July 2009 - Amersham - 24/5 October 2009
Now it's up to you.... Do you want to supercharge your speaking career, speak in exotic locations for high fees and develop a product business or keep going as you have done before. The choice is yours....
Best of Success,
Frank Furness
P.S.- P. S. Please book soon as places are limited and once the bootcamp is filled, the next one will only happen a year later.
Contact Us - Affiliates - Links - Sales Resources - Articles - Free e-books and software Business Links
© 2007 Frank Furness - All Rights Reserved


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Acer Predator G1-710-70011 Gaming Desktop PC - Intel® Core i7-7700, 3.60GHz Processor, 16GB RAM, 2TB HDD + 512GB SSD, DVDRW, NVIDIA® GeForce® GTX 1070 Graphics, Windows 10 Home - UD.P01AA.571Item#: 40629887 | Model#: UD.P01AA.571
- Price:
- $1,809.99
$6.99 Shipping
- Shipping:
- Temporarily out of stock. Order today and we'll deliver when available. (Details)
Additional Resources
Acer Predator G1-710-70011 Desktop - UD.P01AA.571 Product Details
Acer Predator G1-710-70011 Gaming Desktop PC
Equip yourself with a powerful gaming desktop computer for the extensive gaming marathon with the Acer Predator G1-710-70011 Gaming Desktop PC. The Intel® Core i7-7700, Quad-core Processor 3.6GHz lets you experience superb gaming performance without interruptions. NVIDIA® GeForce® GTX 1070 Graphics enhances your gaming sessions by adding realism by producing stunning graphics to your gameplay. 16GB DDR4 RAM memory maintains a good system speed, prevents lags and slowdowns during intense marathon gaming sessions. 2TB HDD + 512GB SSD helps you store tons of games, applications and media files. The Windows 10 Home 64-bit operating systems helps you experience a user-friendly interface. Get yourself a powerful processor that packs in all the advanced major features you need. The Acer Predator G1-710-70011 Gaming Desktop PC has an Intel® Core i7-7700, 3.6GHz Quad-core Processor that will give you the speed and reliability for all you computing tasks. This Desktop is equipped with 16GB RAM memory that you can count on to optimize your computers performance and allow you to run heavy programs without difficulty. It has 2TB HDD + 512GB SSD storage capacity which will give you an enormous storage platter for all your important files. Enter the exciting world of gaming with the Acer Predator G1-710-70011 Gaming Desktop PC
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Apple Products Apps Automotive Babies & Kids Bags & Luggage Beauty & Personal Care Blu-ray, DVD, MP3 Books & Magazines Cameras Cell Phones & Tablets Clothing & Accessories Computer Components Computers Electronics (other) Food & Restaurants
$34.95 Kids Table and Chairs Play Set Toddler Child Toy Activity Furniture In-Outdoor +FS
Ebay offers Kids Table and Chairs Play Set Toddler Child Toy Activity Furniture In-Outdoor for $34.95 (orig. $69.95) with free shipping.
Product Description:
OxGord presents our Kids table and chair set. Made with durable BPA free plastics this colorful Table and Chairs Set is perfect for playrooms, classrooms, daycares, Sunday schools, or kids rooms. 5 bright colors are ideal for both girls and boys rooms. Use as a game table, Lego table or for your children's favorite toys, puzzles, or coloring books. Also perfect as a kids desk or children's lunch table. Durable plastics are tough enough to stand up to the wear and tear of kids but light enough for them to easily move on their own. Easy to clean surface won't mark or scuff walls or floors. 10 inch seat height is perfect for children 3 and up and the 20x20 desk is just the right size for play without taking up too much room in your home. Table and Chair set can be disassembled for easy storage or travel. A must have for daycares and nursery's. Also Ideal for waiting rooms, and doctors offices, any space where children need a play place. Promote your child's imagination while providing them a safe and colorful place to learn and play with this fun and affordable table and chairs set. Table and Chairs come disassembled and can be put together without tools or hardware in a snap.
Thanks Maggie
Category: Babies & Kids
Posted on 09 September 2016, 12:37 PM


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A milestone, of sortsFeb. 12th, 2012 03:20 pm
There are now 365 photographs in my Project 365 set. Sadly, this doesn't actually mean that the goal is complete, because I've missed days here and there and had to start over. My current run started on 30 November.
Onwards and upwards, eh?


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African-American man standing in a field of Austrian winter peas, Windsor, North Carolina, April 4, 1939
Item information
- Title:
- African-American man standing in a field of Austrian winter peas, Windsor, North Carolina, April 4, 1939
- Topics:
- Original Format:
Black-and-white print (photograph)
- Extent:
- 9 x 15 cm
- Item identifier:
- 0050351
- Created Date:
- Description:
- Peason the left were inoculated with commercial cultire. Peas on the right were not inoculated
- Genre:
Portrait photographs
- Names:
North Carolina Cooperative Extension Service
more info on North Carolina Cooperative Extension Service
- Location:
Bertie County (N.C.)
- Digital Project:
University Archives Photographs
Living Off the Land
Source information
- Repository:
- Special Collections Research Center at NCSU Libraries
- Collection:
- Crop and soil science. Grain and legume crops (UA023.007.083) held by Special Collections Research Center at NCSU Libraries
- Note field:
- Not all materials from the physical collection may have been scanned. Images may have been enhanced for web access.
- Rights:
- For questions regarding copyright and permissions, please refer to our Reproduction, Use, Citation, and Copyright page (http://d.lib.ncsu.edu/collections/about).
- Funding:
- Digitization of this material was made possible, in part, through the support of the North Carolina Farm Bureau, the North Carolina Tobacco Foundation, and the North Carolina Tobacco Research Commission.


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14 NBA DFS Podcast for Dec.
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Clarksburg, MA16 Belmar Dr
Est. Mortgage: $/month
MLS Number 220047
- Listings Residential
- Address 16 Belmar Dr
- Bedrooms 3
- Bathroom 2
- Square Feet 2316
Custom Ranch with finished basement and fantastic private yard with huge upper and lower deck and inground pool. Perfect for entertaining! Over 2300 sqft that includes the 750sqft of finished basement area with built in bar area! Beautiful Oak hardwood floors though out dining, living, and all bedrooms. Nice Kitchen, updated in 2008 with White cabintry and Quartz counters! Desirable Clarksburg neighborhood! This beauty won't last long!
Listed by Sue Colvin of Monarch Realty Inc
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|Down Payment
|Interest Rate
Est. Mortgage: $/month
Federal 30-year interest rate: 4.03% last updated on 2017-07-13
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