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Moreover order doxycycline 200mg with visa, NAA changes are sensi- of these amino acids doxycycline 200 mg without a prescription, and has been proposed as a more tive measures of dynamic neuropathologic processes sensitive marker of turnover of the glutamate amino acid (161–163) buy 200mg doxycycline amex, for example, correlating over time with cogni- pool. This has led to speculation that NAA tion of 11 chronic patients and 11 controls, using a 12-cm3 reductions occur as a manifestation of changes in neuronal voxel. They found decreases in NAA peaks in both voxels, volume or in NAA concentrations within a neuron, perhaps with the difference on the right significant at the. Since the report of Nasrallah or a change in the abundance and patterns of neuronal con- et al. It is interesting to note that in various nia have appeared. Most of them have addressed metabolite conditions associated with tissue volume loss and reduced changes, primarily NAA, in the frontal and temporal lobes. Several recent reviews describe these reports in only condition in which NAA concentrations are increased greater detail (143,144,172). Reli- Two other prominent peaks are seen in the long echo ability data for NAA are much superior to those of other time proton spectrum. It is interesting in this regard that all of the MRSI lipid moieties, including glycerophosphocholine, phospho- studies to date that have examined cortical regions in the choline, and phosphatidylcholine. In pathologic conditions frontal and temporal lobes have reported reduced NAA sig- associated with membrane turnover or gliosis (e. These metabolites par- mL voxels in approximately 25 minutes. The technique al- ticipate as energy buffers in many energy-consuming pro- lows for registration of spectroscopic and anatomic images, cesses in the brain, but consistent changes in the CRE signal for reliable and anatomically correct ROI definition, and are generally found only in the presence of tissue loss. The studies At short echo time, several other peaks are observed, in of Bertolino et al. The most studied of these is the myoinositol peak hood-onset patients. Using a similar tech- from glutamate, glutamine, and GABA. Myoinositol is a nical approach in studies of chronic patients, Deicken et al. Chapter 59: Neurochemical and Neuropharmacological Imaging in Schizophrenia 847 TABLE 59. There have been few studies of other peaks in the proton The only imaging study that reported no decrease in gray spectra, and the results have been inconclusive. Because of matter NAA did not regionally parcellate the cortex, and interest in glutamate and cortical function in schizophrenia, reported only an average measure for the entire cortex of several groups have attempted to measure the tGlx peak, the top half of the brain (182). Indeed, this result is consis- using quantitation methods based on a priori knowledge of tent with the findings of Bertolino et al. It is clear from this litera- peaks have generally been negative, and the occasional posi- ture that reduced NAA peaks are found in the frontal and tive result has been inconsistent across studies. The NAA changes have consistently been shown This evidence of relationships between NAA signals and not to correlate with stage of illness, with medication status, connections between neurons led to a series of studies aimed and with length of illness. Moreover, hippocampal and pre- at testing hypotheses about the centrality of prefrontal con- frontal volume do not appear to correlate either. In the nectivity in the pathophysiology of schizophrenia (193). NAA signals selec- suggest that NAA reductions are not the result of a linearly tively in DLPFC were found to predict the availability of progressive pathologic process. They also provide evidence DA receptors in the striatum in patients with schizophrenia, against the notion that chronic untreated psychosis is 'neu- assayed with radionuclide imaging (194). As such, they are unique in data suggested that NAA in DLPFC predicted the steady- vivo evidence of cellular changes probably restricted to neu- state and stimulus-induced responses of dopamine neurons rons in the schizophrenic brain. Similar relationships were not are found in both medicated and unmedicated patients, the found for any other cortical regions or in normal controls, assumption that NAA changes are independent of medical implicating the neuronal pathology associated with the ill- treatment may be incorrect.

Moreover discount 100mg doxycycline otc, this action of lithium signaling pathway cheap 100 mg doxycycline with mastercard, known to be instrumental in normal in the brain and hippocampal cells is apparent only after dorsal–ventral axis development in the Xenopus embryo long-term administration and persists beyond abrupt discon- (108–111) cheap 200 mg doxycycline free shipping. Furthermore, studies in which an embryo ex- tinuation of the drug for an extended period of time, paral- pressing a dominant negative form of GSK-3 was used have leling the clinical time course of the therapeutic effects of demonstrated that myo-inositol can reverse the resulting ab- lithium during initial treatment and discontinuation. Sub- errant axis development in Xenopus, which suggests that sequent studies have discovered that this property of reduc- myo-inositol reversal of dorsalization of the embryonic axis ing the expression of MARCKS in hippocampal cells is by lithium may be mediated, at least in part, by events shared by the anticonvulsant valproic acid, but not by other independent of IMPase inhibition (47). Substrates for GSK- classes of psychotropic agents (96). Additionally, therapeu- 3 in cells include not only glycogen synthase but also - tic concentrations of combined lithium and valproate have catenin and microtubule-associated proteins (MAPs), both induced an additive reduction in MARCKS, also consistent of which have been implicated in cytoskeletal restructuring; with experimental findings that the two drugs workthrough further, -catenin is known to play a role in the expression different mechanisms on the PKC system and the clinical of transcription factors [e. Recent studies in human neuroblastoma cells have demonstrated that valproic acid responses (96). The altered expression of MARCKS further also inhibits GSK-3 , after which levels of -catenin in- supports the role of PI signaling and PKC in the action of crease (112). Thus, GSK-3 may contribute to our under- long-term lithium in the brain and may serve to provide standing of an action for long-term lithium observed in insight regarding a role for neuroplasticity in the long-term events associated with apoptosis and neuroplasticity, as dis- treatment of BPD, as discussed below. A crucial component of cAMP signaling is protein kinase A (PKA), which is a principal mediator of cAMP action in Gene Expression the central nervous system. Long-term lithium treatment has been shown to increase the regulatory and catalytic sub- The clinical data indicating that onset of the therapeutic units of PKA in rat brains, an effect that appears to result effect of lithium requires days to weeks of lag time and that in increased cAMP binding (97). Consistent with a lithium- reversal of the therapeutic effect on discontinuation occurs induced increase in basal cAMP and adenylyl cyclase levels, during a period of weeks to months suggest that the thera- a more recent study has reported that platelets from lithium- peutically relevant action of lithium in the brain involves treated euthymic patients with BPD demonstrated an en- long-term neuroplastic changes mediated by gene regula- hanced basal and the cAMP-stimulated phosphorylation of tion. Evidence has accumulated that lithium can regulate 1146 Neuropsychopharmacology: The Fifth Generation of Progress gene expression via nuclear transcriptional factors. One of neurons, encodes one of the protector proteins that inhibits the immediate early genes, c-fos, works as a master switch apoptosis and cell death under variety of circumstances. Re- of gene regulation through interactions with cis-acting ele- cent studies in rat brain have demonstrated that long-term ments and other transcriptional factors. Lithium has been exposure to lithium or valproate increases the expression of shown to alter the expression of c-fos in various cell systems the polyomavirus enhancer-binding protein 2 gene (113) and in the brain (114–116); however, its effects have (PEBP2B), a regulator of bcl2 expression (133). Subsequent varied depending on brain region, cell type, and time course studies in rat brain have demonstrated an increase in cells examined. In cultured cerebellar granule cells, long-term treat- mon DNA site. Studies in both cell culture and rat brain ment with lithium induces a concentration-dependent de- following long-term lithium exposure in vivo demonstrate crease in p53 and bax (apoptotic genes) mRNA and protein, an enhancement of AP-1 DNA binding activity (99,120). It is of interest that these actions of gene have confirmed a time- and concentration-dependent lithium have been attributed to an enhancement of the PI3 increase in transcriptional activity in the presence of lithium kinase pathway, in which GSK-3 plays a prominent role (121,122). These studies have also noted increases in the (136) (Fig. To what extent this neuroprotective effect protein levels of c-fos, c-jun, and phosphorylated CREB. It may be related to the long-term prophylactic effect of lith- is of interest that phosphorylation of c-jun inhibits DNA ium in stabilizing the course of BPD and the putative role binding, whereas phosphorylation of CREB activates gene of cellular loss in the pathophysiology of affective disorders expression; both are substrates for GSK-3 activity, which remains to be demonstrated (137). However, when AP-1 binding activ- ity was measured following receptor activation, lithium Neuroplasticity and Cytoskeletal treatment attenuated the induced AP-1 DNA binding activ- Remodeling ity (123,124). These seemingly contradictory findings sug- gest that the effect of lithium on gene transcription may Recent studies in a number of laboratories have provided depend on the activity level of the neurons. It has been evidence that long-term lithium treatment may alter molec- suggested that by increasing AP-1 binding activity at the ular substrates underlying neuroplastic changes in brain that basal level, but decreasing it during stimulation, lithium mediate alterations in interneuronal connectivity. As noted can constrain the overall magnitude of fluctuations of gene above, developmental studies in the Xenopus embryo have expression as a function of neuronal activity (125). Valproic recently provided evidence that lithium can act as an inhibi- acid has been shown to have similar effects on the activity tor of GSK-3 , a component of the wnt signaling pathway, of AP-1 (120,122,126), which lends support to the possibil- at concentrations that may be relevant to clinical treatment ity that gene regulation through AP-1 may represent a target (110). Several groups have reported that inhibition of GSK- for mood stabilizers. In addition, carbamazepine has been 3 by lithium reduces phosphorylation of tau protein in shown to inhibit forskolin-induced c-fos gene expression in different cell systems, the effect of which is to enhance the cultured pheochromocytoma (PC12) cells (127).

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These data raise questions regarding the significance of ab- Serotonin (5-HT) is a monoamine neurotransmitter in- normalities in the activity SERT in the pathophysiology of volved in mood and appetite regulation cheap 200 mg doxycycline with visa. In brain order doxycycline 200 mg mastercard, it is syn- depression and the relationship of peripheral and central thesized within the raphe from l-tryptophan buy doxycycline 100 mg otc. There are a number of methodo- self does not cross the blood–brain barrier. Synthesis logic problems inherent in postmortem tissue that may ac- includes an initial conversion to 5-hydroxytryptophan (5 count for differences among studies, including accuracy of HTP) via the enzyme, tryptophan hydroxylase. This system has been 123I- -CIT), that bind selectively to the transporter has al- studied in the brains of suicide victims. In one study, a significant ates reported [3H] phorbol dibutyrate binding to protein difference in binding using SPECT was observed between kinase C in prefrontal cortex was lower in teenage suicide depressed patients and controls (50). More recently they observed that both phos- cant differences were not observed in platelet binding to pholipase C activity and the 1 isozyme protein level were 3H-paroxetine, raising questions regarding whether the decreased of teenage suicide victims (63). Depression per transporter is regulated differently in the two tissues. In contrast Hrdina and associates (64) reported unaltered protein kinase C activity in antidepres- Receptors sant free depressives who suicided, and Coull and colleagues Presynaptic and postsynaptic 5HT receptors have also been (65) reported that phorbol dibutyrate binding sites were studied in depressed patients. Over a dozen serotonin recep- increased in the prefrontal cortex of adults with similar his- tors have been identified, although the possible roles for tories. Age, diagnosis of depression, antidepressant use, and many have not. Two that have attracted most study for the time to collection of brain may play a role in these disparate longest periods are the 5HT1A and 5HT2a types. As with the transporter, multiple studies have ple groups have explored the potential use of pindolol, a investigated 5HT2a binding sites in the platelets of depressed 5HT1a antagonist, to hasten response to antidepressants or patients. An increase in binding sites (B-max) has been re- bring out responses in refractory patients. These studies ported in depressed and suicidal patients with some sugges- have yielded mixed results suggesting that pindolol may tion that increased binding in suicidal patients may be inde- hasten response to antidepressants in milder or first-episode pendent of a diagnosis of major depression (51–53). As with the serotonin transporter, results here activity in suicide victims (66–68). GeneticStudies 5HT2a receptors are found in frontal cortex suggesting a A number of studies have explored the possible role of ge- role in the cognitive aspects of depression. PET ligands have been developed to study 5HT2a activity 18 Long and short forms of the transporter gene appear to be in brain. One study employed [ F]- altanserin and reported relatively common in the general population. An early study a reduction in activity in right posterolateral frontal, orbito- indicated a relationship of the short form with an increased frontal, and anterior cingulate regions in depressives (58). In another study, no differences were found between nonsu- 18 Allelic variation has also been applied to predicting drug icidal depressives and controls using [ F-] setoperone (59). In three studies in Europe and the United States, The exclusion of patients with recent suicidal ideation may homozygotes or heterozygotes for the S-form were reported have played a role in not finding differences between pa- to show sluggish responses to SSRIs (70–72). Studies on effects of antidepressants on was found in a Korean study (73). Clearly, further work is 5HT2a binding using PET have also yielded mix results. More recently, Mann perone binding (60), whereas another recently reported that and colleagues (69) reported that the short form genotype 3 to 4 weeks of desipramine treatment resulted in a signifi- was associated with a diagnosis of major depression but not cant decrease in 5HT2 activity in multiple areas, particularly with suicide or 5HT-transporter binding in postmortem in frontal cortex (61). Depletion Studies 5HT2A receptors are coupled to the phosphoinositide second messenger system. When 5HT2a receptors are acti- Brain concentrations of serotonin are highly dependent on vated by agonists, phosphatidyl inositol 4,5 bisphosphate is circulating levels of tryptophan, which competes with other Chapter 72: Molecular and Cellular Mechanisms in Depression 1043 amino acids for transport into the brain. Charney and Del- theories of depression, DA has been emphasized far less in gado have pioneered in the use of an amino acid cocktail depression in spite of its being widely distributed in brain. In these lite, are decreased in depressives (2,85,86), although some studies, the drink was first administered to subjects who had studies have reported elevated CSF DA, but not HVA levels responded to various antidepressants and who were being (87).

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