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- American Society of Plant Biologists
Plants form shoot meristems in the so-called boundary region, and these meristems are necessary for normal morphogenesis of aerial parts of plants. However, the molecular mechanisms that regulate the formation of shoot meristems are not fully understood. We report here that expression of a chimeric repressor from TCP3 (TCP3SRDX), a member of TEOSINTE BRANCHED1, CYCLOIDEA, and PCF (TCP) transcription factors in Arabidopsis thaliana, resulted in the formation of ectopic shoots on cotyledons and various defects in organ development. Expression of TCP3SRDX induced ectopic expression of boundary-specific genes, namely the CUP-SHAPED COTYLEDON (CUC) genes, and suppressed the expression of miR164, whose product cleaves the transcripts of CUC genes. This abnormal phenotype was substantially reversed on the cuc1 mutant background. By contrast, gain of function of TCP3 suppressed the expression of CUC genes and resulted in the fusion of cotyledons and defects in formation of shoots. The pattern of expression of TCP3 did not overlap with that of the CUC genes. In addition, we found that eight TCPs had functions similar to that of TCP3. Our results demonstrate that the TCP transcription factors play a pivotal role in the control of morphogenesis of shoot organs by negatively regulating the expression of boundary-specific genes.
Meristems are composed of small populations of undifferentiated cells. Plants can regenerate entire organs from shoot meristems via the production of organ primordia. Once organ primordia have been generated, they differentiate into various organs according to their specified fates and cannot return to the meristematic phase during normal development. Lateral organ primordia are established in the peripheral region of a shoot meristem, which is associated with the formation of boundaries that separate organ primordia from the shoot meristem (Aida and Tasaka, 2006a, 2006b). In dicotyledonous plants, a shoot apical meristem (SAM) is formed in the boundary region of two cotyledonary primordia during embryogenesis, and secondary shoot meristems are formed at the boundary of stems and leaves, namely, at leaf axils.
Several factors have been identified as regulators of the formation of shoot meristems in boundary regions. The expression of boundary-specific genes for NAC domain transcription factors, namely, NO APICAL MERISTEM, CUP-SHAPED COTYLEDON (CUC), and CUPULIFORMIS in petunia (Petunia hybrida), Arabidposis thaliana, and Antirrhinum majus, respectively, is necessary for the initiation of formation of shoot meristems (Souer et al., 1996; Aida et al., 1997; Vroemen et al., 2003; Weir et al., 2004). Moreover, loss of expression of two of the three CUC genes in Arabidopsis results in the fusion of cotyledons and defects in the formation of the SAM (Aida et al., 1997; Vroemen et al., 2003). By contrast, ectopic expression of CUC1 enhances the expression of class I KNOTTED1-like homeobox (KNOX) genes and induces the formation of ectopic shoots on the adaxial surface of Arabidopsis cotyledons (Takada et al., 2001; Hibara et al., 2003). This observation indicates that the expression of the CUC1 gene is sufficient for the induction of formation of ectopic shoots on cotyledons. The expression of CUC genes is detected only at the boundaries of embryonic cotyledonary primordia and postembryonic organs, such as leaf axils and floral organs (Aida et al., 1999; Takada et al., 2001; Vroemen et al., 2003; Hibara et al., 2006). Thus, the spatially restricted expression of CUC genes is important for the formation of a shoot meristem at a defined position. Although it has been suggested that the auxin response pathway and microRNA (miRNA) might be involved in the control of the expression of CUC genes (Vernoux et al., 2000; Aida et al., 2002; Furutani et al., 2004; Laufs et al., 2004; Mallory et al., 2004; Baker et al., 2005), the molecular mechanisms that regulate the spatial expression of CUC genes, which are involved in the formation of shoot meristems, remain unknown.
The TEOSINTE BRANCHED1, CYCLOIDEA, and PCF (TCP) family of transcription factors has been reported to play roles in various aspects of plant development (Luo et al., 1996; Doebley et al., 1997; Kosugi and Ohashi, 1997; Cubas et al., 1999). Loss of function of a TCP gene, the CINCINNATA (CIN) gene, in Antirrhinum results in the abnormal curvature of leaves and petals (Nath et al., 2003; Crawford et al., 2004). In addition, suppression of the expression of TCP genes by the ectopic expression of miR319/JAW, which targets TCP2, TCP3, TCP4, TCP10, and TCP24 genes, induced a cin-like phenotype in Arabidopsis (Palatnik et al., 2003). The Arabidopsis genome contains 24 TCP genes, which have been classified into the CYC/TB and PCF subfamilies (see Supplemental Figure 1 online; Cubas, 2000). However, the functional roles of members of the TCP family, including CIN, remain to be clarified.
We developed a gene silencing system, designated chimeric repressor gene-silencing technology (CRES-T), in which a transcription factor fused to the EAR-motif repression domain (SRDX) dominantly represses the transcription of its target genes, even in the presence of endogenous and functionally redundant transcription factors (Hiratsu et al., 2003). Using the CRES-T system, which involves generation of a dominant repressor, we have obtained some insights into the function of TCPs in the regulation of morphogenesis of shoot lateral organs, including the formation of the shoot meristems, via negative control of the expression of boundary-specific genes.
The Chimeric TCP3 Repressor Induces the Formation of Ectopic Shoots
As part of our efforts to identify the biological functions of TCP transcription factors, we applied our CRES-T system to TCP transcription factors because we observed no visible abnormal phenotypic features in either Arabidopsis T-DNA–tagged lines for the TCP genes (data not shown). We converted TCP3, which is phylogenetically close to CIN (see Supplemental Figure 1 online), into a chimeric repressor by fusing it with the SRDX repression domain (Hiratsu et al., 2003), and we expressed the fused gene under the control of the cauliflower mosaic virus (CaMV) 35S promoter (35S:TCP3SRDX; Figure 1A ) in Arabidopsis. Expression of TCP3SRDX induced various morphological abnormalities, which were specifically evident on the surface and at the margins of various organs. Cotyledons of 35S:TCP3SRDX plants were wavy, serrated, and much smaller than those of the wild type, and a number of ectopic shoots were generated on the adaxial side of the cotyledons (Figures 1B to 1E). The abnormal phenotypes of 35S:TCP3SRDX seedlings could be grouped into three classes according to the severity of abnormalities. Seedlings with a mildly abnormal phenotype had epinastic cotyledons with indistinctly differentiated petioles and blades but no ectopic shoots (Figure 1C). Seedlings with a moderate phenotype had cotyledons with serrations at their margins and many ectopic shoots (Figure 1D). The seedlings with a severe phenotype had multiple ectopic shoots on their cotyledons, which resulted in defects in the expansion of the cotyledon (Figures 1E and 2B ). In addition, we often observed ectopic trichomes on the surface of the cotyledons (Figure 2B). Severe defects in the elongation of the main root were also frequent (see Supplemental Figure 2 online), and the pattern of vein formation was severely disrupted in the cotyledons of 35S:TCP3SRDX plants (Figures 1F and 1G). Because the defects in the cotyledons were also evident in mature embryos (Figures 1H and 1I), we postulated that the abnormal development of the cotyledons in 35S:TCP3SRDX plants occurred during embryogenesis.
The seedlings of 35S:TCP3SRDX plants with a mild phenotype grew relatively normally, but their rosette leaves were wavy and serrated (Figures 1J and 1K), resembling those of jaw-D plants (Palatnik et al., 2003). The 35S:TCP3SRDX plants with a moderate or severe phenotype often failed to grow when transferred to soil, and those that developed to rosette plants usually had a bushy phenotype, which was probably due to the presence of ectopic shoots and suggested that the ectopic shoots induced by TCP3SRDX had functional meristems (Figures 1L and 1M). Ectopic shoots were occasionally generated, but at low frequency, on the leaves of 35S:TCP3SRDX plants (data not shown). The morphology of the floral organs of 35S:TCP3SRDX plants was severely abnormal (Figures 1N to 1U). The sepals and petals were wavy and serrated (Figures 1Q and 1S), and the stamens often had clumped outgrowths of cells on their anthers (Figure 1U). Carpels of 35S:TCP3SRDX plants appeared normal, but the surface of siliques was crinkled and the siliques were significantly shorter than those of the wild type (Figure 1V). These results indicated that TCP3SRDX was able to induce abnormal development in various organs, regardless of their identity.
Observations by scanning electron microscopy revealed that wild-type cells in the epidermis and the marginal region were organized in a specific pavement-like pattern and were rod-shaped, respectively, whereas the cells of 35S:TCP3SRDX plants were rounded both in the epidermis and in the marginal regions, with features of undifferentiated cell clusters (Figures 2C to 2F; Donnelly et al., 1999; Ori et al., 2000). Because the differentiation of cells is regulated by their relative position within an organ (Donnelly et al., 1999; Ori et al., 2000), these observations suggested that the epidermal cells of 35S:TCP3SRDX plants do not undergo position-dependent differentiation.
A transient expression assay revealed that TCP3 had transactivation activity and that TCP3SRDX acted as a repressor in Arabidopsis leaves (see Supplemental Figure 3 online). In addition, TCP3mSRDX, in which the encoded amino acid sequence of the SRDX repression domain was mutated (mSRDX; Hiratsu et al., 2004), had no repressive activity (see Supplemental Figure 3 online). We confirmed that the expression of 35S:TCP3mSRDX in transgenic Arabidopsis was unable to induce morphological defects in cotyledons and in leaves (data not shown). These results indicated that the phenotype of 35S:TCP3SRDX plants was induced by the repressive activity of TCP3SRDX and not by some nonspecific negative effect(s), such as squelching (Cahill et al., 1994).
We expressed TCP3SRDX under the control of the 5′-upstream region of the TCP3 gene (ProTCP3:TCPS3RDX), instead of the CaMV 35S promoter, to examine the activity of TCP3SRDX in a condition similar to that of native TCP3. We found that ProTCP3:TCPS3RDX plants had the same phenotype as that of 35S:TCP3SRDX plants, although the frequency of the moderate and severer phenotypes was lower than in the case of 35S:TCP3SRDX plants (Figure 3 ). ProTCP3:TCPS3RDX plants with a mild or a moderate phenotype had leaves sepals, petals, and siliques with wavey surfaces and serrated margins (Figures 3C to 3G). These results indicate that TCP3SRDX, which is present at a concentration more similar to that of the corresponding native transcription factor, could induce the defective phenotype.
TCP3 Regulates the Expression of Boundary-Specific Genes
The 35S:TCP3SRDX plants were morphologically similar to transgenic plants that expressed CUC1 (35S:CUC1) ectopically with respect to the formation of ectopic shoots on the adaxial surface of cotyledons, wavy margins, irregular formation of vasculature, and the undifferentiated rounded shape of epidermal cells (Takada et al., 2001; Hibara et al., 2003). To analyze the effects of TCP3 on the regulation of the spatial expression of CUC1, we introduced TCP3SRDX into an enhancer trap line of CUC1, namely, M0223 (Cary et al., 2002). In M0223 plants, we only detected the promoter activity, as displayed by the fluorescence of green fluorescent protein, in the boundary region between two cotyledons (Figure 4A ). By contrast, the region in which CUC1 was expressed had expanded broadly in cotyledons of M0223 plants that had been transformed with the TCP3SRDX construct (Figure 4B). Resembling results for the CUC1 gene, the areas of expression of two other boundary-specific genes, CUC3 and LATERAL ORGAN BOUNDARIES (LOB) (Shuai et al., 2002; Vroemen et al., 2003), as represented by signals due to β-glucuronidase (GUS) in the enhancer trap WET368 and ET22 lines, respectively, were broadly expanded in the cotyledons of the respective enhancer trap lines, when TCP3SRDX was expressed after it had been introduced by transformation (Figure 4C to 4F). In addition, inappropriate expression of CUC3 was also apparent in mature leaves and sepals of the WET368 line in association with the expression of TCP3SRDX, while no expression of CUC3 was evident in leaves and sepals in the absence of the TCP3SRDX transgene (Figures 4G to 4K). Cells in leaves in which the CUC3 gene was ectopically expressed were often rounded (Figure 4I), suggesting that the expression of the boundary-specific genes induced an undifferentiated state in these cells. Analysis of the expression of transcripts revealed that the boundary-specific genes CUC1, CUC3, CYP78A5, LOB, LATERAL SUPPRESSOR, and BLADE ON PETIOLE1 (BOP1) (Aida et al., 1997; Zondlo and Irish, 1999; Shuai et al., 2002; Greb et al., 2003; Vroemen et al., 2003; Ha et al., 2004) were expressed in leaves of 35S:TCP3SRDX plants, whereas these genes were not expressed in leaves of wild-type plants (Figure 4N). Similarly, the CUC1, CUC3, and LOB genes were also expressed ectopically in the cotyledons of ProTCP3:TCP3SRDX plants (see Supplemental Figure 4 online). These results indicate that TCP3SRDX induced the ectopic expression of a variety of boundary-specific genes in a variety of organs.
In addition to its effect on boundary-specific genes, we examined the effect of TCPSRDX on the expression of class I KNOX genes, namely, KNAT1, KNAT2, and SHOOT MERISTEMLESS (STM), which is required for the formation of a functional meristem (Chuck et al., 1996; Ori et al., 2000; Hake et al., 2004). The promoter activity of KNAT1 was detected only in the SAM of ProKNAT1:GUS plants, while the region in which KNAT1 was expressed was much more extensive in the presence of TCP3SRDX (Figures 4L and 4M). In addition, analysis by RT-PCR showed that KNAT1, KNAT2, and STM were expressed ectopically in leaves of 35S:TCP3SRDX plants (Figure 4N). Since the product of the CUC1 gene is a positive regulator of the expression of class I KNOX genes (Takada et al., 2001; Hibara et al., 2003), it seems likely that enhanced expression of boundary-specific genes, including the CUC1 gene, in response to TCP3SRDX induced the inappropriate expression of the class I KNOX genes in 35S:TCP3SRDX plants.
The expression of CUC genes is negatively regulated by miR164 (Laufs et al., 2004; Mallory et al., 2004; Baker et al., 2005). To investigate whether TCP3 is involved in the accumulation of miR164, we examined levels of miR164 in 35S:TCP3SRDX plants. RNA gel blot analysis revealed a significant reduction in the accumulation of miR164 in 35S:TCP3SRDX plants (Figure 5 ), suggesting the involvement of TCP3 in the regulation of the accumulation of this miRNA.
Loss of CUC Activity Suppresses the Function of the Chimeric TCP3 Repressor
To confirm that the abnormal phenotype of 35S:TCP3SRDX plants was due to the inappropriate expression of boundary-specific genes, we expressed TCP3SRDX in plants with a loss-of-function mutation in a boundary-specific gene. When TCP3SRDX was expressed on the cuc1 mutant background, the defective phenotype of cotyledons of 35S:TCP3SRDX plants was suppressed to a considerable extent and most of the seedlings had normal cotyledons with a flat surface and smooth margins, with no ectopic shoots (Figures 6A and 6B ). Such recovery was observed similarly on the cuc2 mutant background, but the frequency of recovery was lower than on the cuc1 mutant background (Figure 6C). By contrast, no recovery was observed on the bop1-4 mutant background (data not shown). These results indicated that the abnormal phenotype of 35S:TCP3SRDX plants was most likely due to the inappropriate expression of the CUC genes.
Gain of Function of TCP3 Suppresses the Formation of Shoot Meristems
Transgenic plants that expressed TCP3 ectopically (35S:TCP3) had no visible abnormalities, probably as a result of the activity of miR319/JAW (data not shown). Therefore, we expressed a mutant form of TCP3 (mTCP3) in which the target site of miR319/JAW had been replaced by a nontarget sequence, without any change in the encoded amino acid sequence, as described previously in the case of TCP2 and TCP4 (Palatnik et al., 2003). We found that 35S:mTCP3 induced the fusion of cotyledons and defects in the formation of shoots, in addition to enhanced elongation of hypocotyls (Figures 7A to 7C ). This phenotype was somewhat similar to that of the cuc1 cuc2 double mutant. Similar fusion of cotyledons was also observed in 35S:mTCP2 and 35S:mTCP4 plants to varying degrees (Palatnik et al., 2003; see Supplemental Figure 5 online). In seedlings of these plants, the expression of CUC1 and CUC3 was significantly suppressed or undetectable (Figure 7D; see Supplemental Figure 5 online). Analysis by RT-PCR confirmed that the level of the expression of these CUC genes was clearly reduced in 35S:mTCP3 plants (data not shown). These results demonstrated that TCP3, in addition to TCP2 and TCP4, can suppress the expression of CUC genes.
Redundant Functions of the Members of the TCP Family
As compared with transgenic plants that expressed TCP3SRDX, we found that tcp3-1 plants, namely, the TCP3 T-DNA–tagged homozygous line (CS855978), and transgenic plants that expressed the double-stranded RNA for RNA interference (RNAi) of the TCP3 gene had basically normal cotyledons (see Supplemental Figure 6 online). By contrast, ectopic expression of a genomic DNA fragment that encoded miR319/JAW, which should cleave transcripts of the TCP2, TCP3, TCP4, TCP10, and TCP24 genes, resulted in cotyledons that resembled those of 35S:TCP3SRDX plants with the mild phenotype (see Supplemental Figure 6 online), in addition to an effect on leaf phenotype, as reported previously (Palatnik et al., 2003). These analyses suggest that suppression of the expression of five TCP genes might be required for induction of cotyledons with the abnormal phenotype because of the functional redundancy of TCP genes.
We produced seven lines of transgenic Arabidopsis plants that expressed individual chimeric repressors derived from each of seven other TCPs in the CYC/TB subfamily, namely, TCP2, TCP4, TCP5, TCP10, TCP13, TCP17, and TCP24. We found that all seven chimeric repressors induced phenotypes similar to that induced by TCP3SRDX, although the severity of the phenotypes differed even when gene expression was driven by the CaMV 35S promoter (see Supplemental Figures 6A and 6B online). In these transgenic plants, we confirmed that the boundary-specific genes were expressed ectopically in cotyledons and leaves, as they had been also in 35S:TCP3SRDX plants (see Supplemental Figures 7A and 7B online). Moreover, expression of TCP10SRDX driven by its homologous promoter (ProTCP10:TCP10SRDX) induced defects in cotyledons and the ectopic expression of boundary-specific genes similar to those observed when transcription was driven by the CaMV 35S promoter (see Supplemental Figure 8 online). These observations suggested that the TCP transcription factors examined in this study have similar molecular functions and are possible regulators of the expression of boundary-specific genes.
The Patterns of Expression of the TCP Genes
In situ hybridization revealed that the TCP3 transcript was localized in the distal and middle regions of cotyledons of torpedo, bending cotyledon, and mature embryos, but no signals were detected in the presumptive SAM and the boundary region during embryogenesis (Figures 8A to 8E ). Resembling the TCP3 transcript, the TCP10 transcript was also detected in regions of the developing embryo except the region of the presumptive SAM and the boundary region (Figures 8F to 8I). In contrast with the TCP3 transcript, however, the TCP10 transcript was detectable in the cotyledons of heart-stage embryos (Figure 8F). Since expression of the TCP4 gene is detectable in embryonic cotyledons (Palatnik et al., 2003), these three TCP genes have similar patterns of expression in cotyledons.
We also examined the patterns of expression of TCP genes using a GUS reporter gene that was fused with the 5′-upstream region of each respective TCP gene (ProTCP:GUS). The promoter activity of the TCP3 gene, as represented by the GUS activity due to the ProTCP3:GUS gene, was strong in the vascular regions of the cotyledons and hypocotyl in torpedo and bending cotyledon embryos, as well as in developing seedlings (Figures 8J to 8N). The difference between the patterns of expression of TCP3 during early embryogenesis obtained with ProTCP3:GUS and by in situ analysis might have been due to posttranscriptional suppression by the miR319/JAW-mediated cleavage of the TCP3 transcript or to transcriptional regulation via sequences beyond the promoter region used in this study. The promoters of the TCP2, TCP4, TCP5, TCP10, TCP13, TCP17, and TCP24 genes were active in cotyledons and, in particular, in the vascular region, as was the case for the promoter of the TCP3 gene, even though each gene had a somewhat different pattern of expression (see Supplemental Figure 9 online). Analysis by RT-PCR confirmed that these eight TCP genes are expressed differentially in various organs in an overlapping manner, although no signal for TCP5 was detected in roots (Figure 8O).
In this study, we found that the expression of a chimeric TCP3 repressor induced the ectopic expression of boundary-specific genes, with the resultant formation of ectopic shoots, while overexpression of mTCP3 suppressed the expression of such genes, with resultant inhibition of the formation of shoots and of the separation of cotyledons. Our results demonstrate that TCP3 plays a pivotal role in the control of morphogenesis of shoot lateral organs via the negative control of the expression of boundary-specific genes, namely, the CUC genes, which control the initiation of the shoot meristem and the morphogenesis of organ boundaries (Aida et al., 1997). Given that TCP3 has transactivational activity, we can postulate that TCP3 might activate some unidentified factors that suppress the transcription of the CUC genes. The proposed function of TCP3 is consistent with the finding that the pattern of expression of the TCP3 gene does not overlap with that of CUC genes.
The expression of the CUC genes is regulated at the transcriptional level but is also regulated negatively by miR164 (Laufs et al., 2004; Mallory et al., 2004; Baker et al., 2005). miR164 is encoded by three genes, miR164A, miR164B, and miR164C, and it accumulates in various tissues, which include seedlings, leaves, and floral organs (Laufs et al., 2004; Mallory et al., 2004; Baker et al., 2005). The suppression of expression of miR164 in a T-DNA–tagged line for the miR164A gene deepened the serration of leaf margin (Nikovics et al., 2006), and this phenotype was somewhat similar to that of 35S:TCP3SRDX plants. Thus, control of the accumulation of miR164 might be important for appropriate development of leaves. We showed in this report that the level of miR164 was significantly reduced in 35S:TCP3SRDX plants (Figure 5). It is possible that suppression of the expression of the miR164 gene is also involved in the ectopic expression of CUC genes in 35S:TCP3SRDX plants. However, considering that mutation of the miR164A gene did not induce ectopic shoots, we postulate that TCP3 might control the expression of the CUC genes at both the transcriptional and posttranscriptional levels.
Our results suggest that elimination from cotyledons of the activities of proteins encoded by the CUC and other boundary-specific genes might be required for the normal development of shoot lateral organs. Since the CUC genes and several boundary-specific genes appear to control morphogenesis at boundaries and the formation of shoot meristems (Aida and Tasaka, 2006a, 2006b), inappropriate expression of these genes might prevent the specific differentiation of cells that is necessary for the formation of smooth surfaces and margins of organs. Indeed, ectopic expression of the CUC genes and other boundary-specific genes has been reported to disturb the normal development of the respective transgenic plants (Zondlo and Irish, 1999; Takada et al., 2001; Shuai et al., 2002; Hibara et al., 2003; Laufs et al., 2004). The abnormal distribution of or response to auxin induces the inappropriate expression of CUC genes in cotyledons and lateral organ primordia, preventing organ development (Vernoux et al., 2000; Furutani et al., 2004; Treml et al., 2005). In addition, the abnormal radial information generated by inappropriate expression of PINHEAD in the peripheral region of an embryo induces ectopic expression of CUC2 and the formation of shoot meristems on cotyledons (Newman et al., 2002). Thus, the spatial regulation of the expression of the CUC genes and other boundary-specific genes might be a critical determinant in the control of morphogenesis of shoot lateral organs. TCPs appear to be novel factors that are involved in this regulation.
By contrast, it appears that initiation of the formation of shoot meristems requires suppression of the activities of TCPs in the boundary region and the SAM. The accumulation of transcripts of the TCP3, TCP4, and TCP10 genes and the activities of the respective promoters were not evident either in the SAM or at cotyledonary boundaries (Figure 8; Palatnik et al., 2003). In addition, we found that ectopic expression of mTCP3, in which the target site of miR319/JAW had been mutated, inhibited formation of shoots, probably as a result of the expression of TCP3 in the boundary region. Consistent with these observations, the precursor to miR319/JAW is abundant in the shoot apex, which includes the SAM (Palatnik et al., 2003). Thus, the miR319/JAW-dependent cleavage of TCP transcripts might be required for the formation of the shoot meristem. Our results suggest that suppression of the activities of TCPs in the boundary region might be controlled at both the transcriptional and the posttranscriptional level.
We showed that eight TCPs have similar molecular functions by examining the effects of the chimeric repressors derived from each one, and we found that the corresponding TCP genes were expressed in an overlapping manner in various tissues. The TCP3, TCP4, and TCP10 genes have similar patterns of expression in cotyledons during embryogenesis, and seven of the eight TCP genes are expressed during the early development of flowers (Figure 8; Palatnik et al., 2003; Wellmer et al., 2006). In addition, the functional redundancy of these TCPs was apparent from the observation that neither T-DNA–tagged lines nor RNAi for TCP3 induced an informative phenotype in Arabidopsis. Although 35S:miR319/JAW plants have mildly abnormal cotyledons, the functions of the TCP5, TCP13, and TCP17 genes, whose transcripts lack a target sequence for miR319/JAW, might prevent formation of more severely abnormal cotyledons, with, for example, ectopic shoots. These observations suggest that the TCP transcription factors of Arabidopsis act redundantly to regulate the spatial expression of boundary-specific genes and to control the morphogenesis of shoot lateral organs. Arabidopsis appears to have a greater number of redundant TCP genes than does Antirrhinum because disruption of the CIN gene alone is sufficient to induce abnormal leaves in Antirrhinum (Nath et al., 2003).
Each TCP gene had a somewhat different pattern of expression, and each chimeric TCP repressor induced an abnormal phenotype with a different degree of severity, even when transcription of its gene was driven by the CaMV 35S promoter. The chimeric repressors derived from TCP3, TCP4, TCP5, and TCP10 induced the severe phenotype at higher frequency, perhaps because of higher activities of DNA binding and transactivation. In addition, the activities of the promoter regions of the TCP3, TCP4, TCP5, and TCP10 genes used in this study were more prominent in cotyledons. These genes might make a larger contribution to the development of cotyledons than do other TCP genes. Thus, the TCP transcription factors examined in this study appear to function similarly as regulators of the expression of boundary-specific genes, but the spatial activity of each TCP seems to be regulated differently.
Because of the close relationships among TCPs, in terms of phylogeny, and their functional similarities, we have grouped these eight TCPs as a subfamily of CIN-like TCPs. TCP genes are conserved in monocots, as well as in dicots, and there are 23 TCP genes in the rice (Oryza sativa) genome, just as there are in the Arabidopsis genome. Ten of these rice genes, which include genes for class II PCFs (Kosugi and Ohashi, 2002), can be considered to be CIN-like TCP genes. We found that five of the CIN-like TCP genes in rice have a target sequence for miR319/JAW in addition to those mentioned in a report by Palatnik et al. (2003). Thus, it is likely that not only the family of TCP genes but also the mechanism for regulation of the expression of genes for TCPs by the miR319/JAW gene evolved prior to the divergence of monocots and dicots. It has been suggested that CUC genes might be involved in the formation of the SAM in monocots (Zimmermann and Werr, 2005). Thus, the acquisition of a mechanism for regulation of the spatial activities of CUCs by TCPs might have been a key event in the evolution of plants.
In this study, we showed that TCP transcription factors are critical for the morphogenesis of shoot lateral organs. Extending the previous reports that TCPs are involved in the control of the development of leaves (Nath et al., 2003; Palatnik et al., 2003), we showed that TCPs are involved in the formation of the shoot meristem and the development of shoot lateral organs. When TCPs fail to suppress the expression of the CUC genes, ectopic expression of CUC genes results in the generation of multiple shoot meristems and inhibition of organ growth. Plant cells have extraordinary totipotency and have the ability to generate new organs continuously from shoot meristems and to form shoot meristems at organ boundaries throughout their life cycle (Weigel and Jurgens, 2002; Willemsen and Scheres, 2004; Schmitz and Theres, 2005; Aida and Tasaka, 2006a, 2006b). This process requires the coordination of the functions of the shoot meristem with those of the differentiating cells of a given organ. Thus, the TCP-dependent regulation of the expression of CUC genes, which controls morphogenesis of the shoot lateral organs and regulates formation of the shoot meristem, is of critical importance to plant development.
Construction of Plasmids
The protein-coding regions of TCP genes were amplified from genomic DNA or from cDNAs provided by the RIKEN Bio Resource Center (BRC; Seki et al., 1998, 2002) with appropriate primers, as shown in Supplemental Table 1 online. The chimeric 35S:TCPSRDX gene was constructed as described previously (Mitsuda et al., 2006). The mSRDX, mTCP2, mTCP3, and mTCP4 genes were generated by site-directed mutagenesis with appropriate primers. The GUS reporter genes under the control of the promoters of individual TCP genes were constructed using 5′-upstream regions relative to sites of initiation of translation of the respective TCP genes (TCP2, 2978 bp; TCP3, 2593 bp; TCP4, 3027 bp; TCP5, 2879 bp; TCP10, 1054 bp; TCP13, 2863 bp; TCP17, 2276 bp; and TCP24, 2776 bp). The GUS genes of ProTCP3:GUS and ProTCP10:GUS were replaced by TCP3SRDX and TCP10SRDX for the construction of ProTCP3:TCP3SRDX and ProTCP10:TCP10SRDX, respectively. The construct for RNAi of TCP3 was generated from the 458-bp region at the 3′ end of the gene (nucleotides 715 to 1173) and pHELLSGATE8 (Wesley et al., 2001).
Plant Materials and Transformation
Arabidopsis thaliana ecotype Col-0 was used throughout this study unless otherwise indicated. Growth conditions and the strategy for transformation of Arabidopsis were described previously (Mitsuda et al., 2006). The exogenous expression of each transgene in transgenic plants was confirmed by RT-PCR with appropriate primers (see Supplemental Figure 10 and Supplemental Table 1 online). The tcp3-1 line had a T-DNA tag in the protein-coding region of the TCP3 gene at a site 261 bp from the site of initiation of translation.
Light and Scanning Electron Microscopy
Analysis of promoter activities using the GUS reporter gene was performed with T1 or T2 transgenic lines as described previously (Mitsuda et al., 2005). Light microscopy and fluorescence microscopy for detection of green fluorescent protein were performed with the Axioskop2 plus system (Carl Zeiss). Plant tissues were rendered transparent for the observations of vasculature as described previously (Aida et al., 1997). For scanning electron microscopy, samples were prepared and analyzed as described previously (Mitsuda et al., 2005).
Isolation of RNA and RT-PCR
Total RNA was isolated from tissues with Trizol as described previously (Fujimoto et al., 2000). For analysis by RT-PCR, aliquots of 50 ng of total RNA were subjected to first-strand cDNA synthesis (Hiratsu et al., 2003). PCR was performed with gene-specific primers (see Supplemental Table 1 online) for 30 to 39 cycles.
Small RNA was prepared with a mirVana miRNA isolation kit (Ambion) from 3-week-old plants. For detection of miR164, aliquots of 40 μg of small RNA were fractionated on a 15% polyacrylamide gel that contained 7 M urea, blotted onto a nylon membrane, and allowed to hybridize to the 32P-labeled nucleotide probe in ULTRAhyb-Oligo solution (Ambion). The synthetic RNA corresponding to the sense strand of miR164 was used as a positive control for hybridization. The intensity of signals was quantified with ImageQuant (Molecular Dynamics).
In Situ Hybridization
Preparation of samples and in situ hybridization were performed as described previously (Furutani et al., 2004). DNA fragments corresponding to positions 48 to 1176 from the site of initiation of translation of TCP3 and to the full-length coding sequence of TCP10 were used as templates for probes, respectively.
Assays of Transient Gene Expression
The coding sequences of TCP3, TCP3SRDX, and TCP3mSRDX were fused separately to that for GAL4DB (Ohta et al., 2000), and assays of transient gene expression were performed as described previously (Fujimoto et al., 2000). We used 0.8 μg of reporter plasmid and 0.6 μg of effector plasmid for each bombardment. For normalization of the activity of the reporter gene, we used 0.8 μg of a reference plasmid, pPTRL (Fujimoto et al., 2000). After bombardment, samples were incubated for 16 h in darkness, and then luciferase activity was quantified.
Accession Numbers
The Arabidopsis Genome Initiative (http://www.arabidopsis.org) identifiers for the genes and gene products were designated as follows: TCP2, At4g18390; TCP3, At1g53230; TCP4, At3g15030; TCP5, At5g60970; TCP10, At2g31070; TCP13, At3g02150; TCP17, At5g08070; and TCP24, At1g30210.
Supplemental Data
The following materials are available in the online version of this article.
Supplemental Table 1. Primers Used in This Study.
Supplemental Figure 1. Phylogeny of TCPs in the CYC/TB Subfamily.
Supplemental Figure 2. Inhibition of Elongation of the Main Root by the Chimeric TCP Repressor.
Supplemental Figure 3. Transient Expression Assay for TCP3.
Supplemental Figure 4. Expression of Boundary-Specific Genes in ProTCP3:TCP3SRDX Plants.
Supplemental Figure 5. Phenotype Induced by the Ectopic Expression of the mTCP Gene.
Supplemental Figure 6. Phenotype Induced by Suppression of the Expression of the TCP Gene.
Supplemental Figure 7. Phenotypes Induced by Seven Different Chimeric TCP Repressors.
Supplemental Figure 8. Phenotype of ProTCP10:TCP10SRDX Plants.
Supplemental Figure 9. Analysis of the Promoter Activities of TCP Genes in Young Seedlings.
Supplemental Figure 10. Expression of the TCP3SRDX Gene in 35S:TCP3SRDX Plants.
We thank the ABRC for seeds of tcp3-1 (CS855978), ET-22, and ProKNAT1:GUS plants; C. de Vries for seeds of WET368 plants; C.M. Ha for seeds of bop1-4 plants; RIKEN BRC for cDNA clones for TCP genes; and P.M. Waterhouse for pHELLSGATE8. We thank Akita Prefectural University for sequencing plasmids and S. Miyamura and A. Iwase for scanning electron microscopy analysis. We also thank K. Hiratsu, N. Mitsuda, and M. Aida for helpful discussions and K. Yamaguchi, A. Kushida, N. Kawanami, and Y. Takiguchi for their skilled technical assistance.
The author responsible for distribution of materials integral to the findings presented in this article in accordance with the policy described in the Instructions for Authors (www.plantcell.org) is: Masaru Ohme-Takagi (m-takagi{at}aist.go.jp).
↵[OA] Open Access articles can be viewed online without a subscription.
↵[W] Online version contains Web-only data.
- Received June 10, 2006.
- Revised December 21, 2006.
- Accepted January 31, 2007.
- Published February 16, 2006.


text:
Ian R. Dunnigan
Birth Controller to the Jazz Community
About Me
Born at an early age, I grew up in the countryside around Loch Lomond in Scotland - yes THAT Loch Lomond. The one that has the bonnie bonnie banks. At primary school, I took up descant recorder, to avoid the school choir, and on arriving at secondary school, had a flute thrust into my hand. At 14, I acquired a 1937 Buescher Aristocrat Alto Sax, and that became the favoured instrument.
Leaving school to start my apprenticeship as a Chartered Accountant, I realised that the music and the studies were not going to go together, and regretfully the instruments were packed away, except for the odd session with some friends.
After qualifying, I eventually ended up in London, by way of Liberia in West Africa, (where I picked up a dirt cheap Yamaha Tenor), The Isle of Lewis, and Aberdeen in Scotland.
Musically, I started with Rock and Roll and Rhythm & Blues (the real stuff), 60s pop and Scottish folk music. The odd Jazz album started sneaking in from 16 on, and I also started getting a taste for Classical music, which I never had at school, despite having to play it on Flute.
Now living in London, I can indulge in as much Jazz as I want. London has a thriving scene around the pubs and clubs.
As I seem to have received a number of requests to review albums recently. It is not something I do — I'm just no good at it — Sorry

subset: commoncrawl


text:
He told dispatchers he thought she might have taken too much medication for anxiety or depression.
Officers who responded to the call found several prescription bottles, including Ambien and anti-anxiety medication, that had been filled within 10 days but were empty, according to a police report. It also said there was a handwritten note that indicated she had taken the pills intentionally.
The Kings filed for divorce in April, but Larry King released a statement through his spokesman, Howard Rubenstein, saying "reports that we have fought are absolutely false."
Shawn King is Larry King's seventh wife. The couple has been married since 1997 and has two sons.
The Associated Press contributed to this story.
Your feedback is important to us! Please complete a brief survey so we may continue to improve abc7.com


text:
Companies can see how employees compare in terms of computer activity, application use, active/inactive time and more.
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text:
In honor of the Yahrtzeit yesterday of Rabbi Yosef Menachem Mendel Tenenbaum, longtime principal at United Lubavitcher Yeshivah, we present you with several stories he experienced with the Rebbe, compiled and written by Rabbi Dovid Zaklikowski of Lubavitch Archives.
With the Ramash
When the Rebbe was still known as the Ramash, he would come to the beis midrash library in 770 to consult different seforim. Rabbi Tenenbaum once approached the Rebbe there and asked for his approval for an explanation of a certain concept in Chassidus.
After Rabbi Tenenbaum explained his idea, the Rebbe said, "I hear what you are saying, but I don't hold that way…"
Remember the Days
Once, when the Rebbe was encouraging people, during a farbrengen, to daven with more concentration and sincerity, the Rebbe turned to Rabbi Tenenbaum and said, "Zechor yemos olam…" (Remember the days of old…) (Devarim 32:7).
In Debt for a Miracle
George Landberg was an English teacher at the yeshivah when his wife gave birth to a son. In the hospital, one of the nurses mistakenly placed 70 times the prescribed dose of silver nitrate drops into the newborn's eyes (today the nitrate drops come prepackaged). "The skin near the eyes looked like it was burned," Mr. Landberg recalled. The doctors said that the child would be blind in both eyes.
When Rabbi Tenenbaum gave him a mazel tov on the birth of his son, Mr. Landberg broke down in tears, telling Rabbi Tenenbaum the entire story.
Rabbi Tenenbaum tried to calm him down, wrote down the family members' Hebrew names and told him that he would write to the Rebbe for a bracha. The Rebbe gave his bracha to the child, and over the next few weeks the baby's eyesight improved. "The doctors could not believe that it was possible to change for the better," Mr. Landberg said. "They told me that it was a miracle."
At the time Mr. Landberg was not mitzvah observant had little Jewish knowledge. When Rabbi Tenenbaum heard about the miracle thanks to the Rebbe's bracha, he told Mr. Landberg that it was time for him to pay up his debt. "In merit of the miracle, I want you should take a mitzvah upon yourself," Rabbi Tenenbaum said.
Mr. Landberg agreed to put on tefillin every day.
When Mr. Landberg retold the story, he always said that he hadn't missed a weekday of putting on tefillin, and that his son puts on tefillin, too.
Running to the Bus
At one time, Nostrand and Kingston Avenues were two-way streets. The traffic then on Brooklyn Avenue would go towards Eastern Parkway, and New York Avenue went down to Empire Boulevard. In 1962, when they made Nostrand and Kingston one-way streets, the directions of New York and Brooklyn were swapped, so that every street should go in a different direction.
This change created a new opportunity. If all the children who were going home on the bus from yeshivah would make it there on time, they would be able to see the Rebbe returning from 770 to his house on President Street.
Thus the children would run down the hallways to the bus and pressure the bus driver to leave as soon as possible. When they saw the Rebbe walking down Brooklyn Avenue, the bus would slow down, and all the children would run to the left side and sing "Uforatza." The Rebbe would turn towards the bus and encourage the singing.
Rabbi Tenenbaum did not like the running that was going on. Seeing that it was a daily occurrence, he approached the children and requested that they not run.
However, when they told Rabbi Tenenbaum that they were running so they could see the Rebbe returning from 770, he was very pleased and gladly agreed that they could continue to rush to the bus.
London "Touring"
During a yechidus before Uncle Itzy's wedding in England (on Adar 7, 1975), the Rebbe told Rabbi Tenenbaum, "You will surely visit the mosdos in London."
During the days we were in London, Rabbi Tenenbaum went from one Chabad institution to the next. The family could only persuade him to spend a few hours sightseeing at Buckingham Palace.
Dovid Zaklikowski is a historian and writer, his latest books Learning on the Job (amazon.com/dp/194487500X) and Advice for Life (https://www.amazon.com/dp/1944875018) are available on Amazon.com.


text:
The challenge between perfect technology architecture and fast shifting customer channels
There is a lesson here for Banks. In 2012 just over a year ago, FaceBook went public. At the time the largest doubt was FaceBooks ability to leverage advertising in the mobile space.
Since then FaceBook has turned that corner. It has done that by making mobile important.
But how did FaceBook accomplish this and how did they accomplish this in just over a year.
How Facebook Went Mobile, In Before And After Org Charts
Facebook used to be a website translated to mobile by a tiny team, but over the last two years it's reorganized to make every department in the company mobile-first, as revealed in two new org charts it shared today at a small press conference "Whiteboard Session" at its Menlo Park HQ.
I don't totally understand the org charts TechCrunch shows, but the more important point is that the new organisation has focus on mobile first. This focus has led them to develop their own tools that support their scale.
The new company structure also gives Facebook the resources to build critical mobile development infrastructure. For example, Legnitto showed off xctool, a replacement for Apple's xcodebuild that makes it easier to develop iOS and Mac products. Legnitto said "Apple's tools are good but they're designed for the individual developer. Their tools started to fall over at our scale" referring to its 874 million monthly mobile users and 507 million daily mobile users.
Banks have their own scale issues. They have legacy deposit and loan systems that must interact with new CRM systems then finally with branch, web and mobile platforms for customer access. This is no small feat. Hidden in that system diversity is the disconnect between branch, web and mobile. Bank system development has traditionally focussed on branch first. Gradually web has received priority but you get the picture.
In early 2000's I was a strong advocate that we should build for web first, and let that development improvements flow into the branch. I was lone wolf on that view.
Now today I say the answer ought to be development for mobile first. Going back to the FaceBook example the key is to make development of the next platform the most important. Dedicate the resources there and share the results.
Apple are another example of this. The laptop OS is now picking up design elements from iOS. This is the perfect example of designing for customers and their needs first, but it requires internal shifts that are difficult and awkward. It will require changes that will be strongly resisted by the status quo.
The status quo will pull towards the classic Microsoft / IBM model of building once and presenting out to multiple platforms simultaneously. This approach may be logical but it is not what FaceBook with 50,000 employees did when they turned the company on a dime in 12 months. Maybe its better for the product and the clients to drive for the goal first and sort it out later.
There are lessons to be learned here.
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text:
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text:
Cars, Appliances Could Be Hack Targets
It isn't just PCs that are vulnerable to hack attacks, officials warn. The new list of vulnerable objects includes cars, appliances and electricity meters, Jennifer Valentino-Devries reports.
This transcript has been automatically generated and may not be 100% accurate.
kg ... with more ... Michaels would amount to know is still with namely the talk about is understood PC baking day and you thought you will want my photo like this ... gem um ... and frightening story of love that not only tell what to do and the Seychelles ... these can be countries with the electronics and ... bought some tells about is a warning from dry ... you know it's not as though people are at ... any rate isn't anyone stealing cars actually added the real ... issue here and is that's ... their Sony devices now people don't realize that they have computers and then connected TV Internet and cell phone network computers and that is so small ... this technology is ... so easy that ... it everywhere and that actually means that ... although these devices that are connected now ... in their vulnerabilities as potentially they can be exploited by hackers most of the puppet right time ... this is indeed a covert a notice to my second research has demonstrated how they could on law and stop the complex and is the text messages ... to the cost of on rest and let them get cover of more than at any time and so I mean it ... was on a second tunnel was that was that the US and their head site Hulu a video of ... their cybersecurity company and said the conference in San about cybersecurity and then packing and said they use this demonstration and I think they would say that ... I'm that it is that it represents the Lache sees a car because that's something that everybody understands and if your card in the Ummah that's just ... you know we we kind of threatening to lot of people but ... I'm the real point is that this ad ... was either he's telling you right because of the data center here in Cardon in case he probably had a scan ... yesterday were able to monitor the traffic been screened the end of the car this computer and high ... and the self an hour and then they were able to penance he ... added ... what the commands where they got the parts of eastern things and and they use their own computer day ... again over the Soffa network send messages to this computer that said ... the same thing and see when the Proms that when that was that ... enabled device is better in everything now they're so small and they're really trying to be made cheaply on that day ... he act like a comparable computers that control them ... and ... don't necessarily have good security measures photography they don't ... and to keep the messages that are coming to them so ... that is the problem that can really say no it's not just cars and one other thing you want to use the Evo that Sokol with the beat may we get up ... DoubleLine strong ... exports digits tells about that white wine to consensus on people so this this and seeing people like the Act I determined Korean security and electrical providers ... they can think I had this machinery now these these and gadgets with is that ... you might tell them ... how to be head planes is what DNA allows them to better monitor the plant's ands from only give instructions to make sure that you think they're ... upsetting on and off that every time that sort of thing ... and ... that ... you know again like bit secure the system in the car ... these are connected within cellphone network and they don't have the proper infection and ... somebody could same thing ... for them to get them to ... be a turnoff ... at certain generators improperly and then that could potentially cause of the pack or ... an overseas you you you mention that all nonsecurity on my ... to do with terror threats on the wood with some of them in a bubble about to die don't agree with this ... aam but also well inside the wall ... this could be presumed improvements on the Watford keeper Matt computers now ... form a new Iraq or even of infantry soldiers from ... re signing I think I'm ... I think their concern right now is to relieve water with that ... and port infrastructure ... at this price you know the folks that I've talked to thinking of this as a concern for eye for them and these are in heaven one emphasize that these are vulnerabilities and ... United tied to anybody he said that this was being attacked currently and ... the families and that is to say that honesty ... in the country so scared that their cards in a day in and still no sign of how to something like Opana says it killed a security new Brideau computed generally you read things that we just don't read through the palate with Weibo and always ready with all my point is the full level that would open up the ... problems there so many machines it's difficult to get security patches at the end of the vulnerability is the ... sense you know what what what turned out


text:
Balls: Motorcyclist Lays Head On Track In High Speed Turn
This is an older video of a motorcyclist laying his head on the track in a high-speed turn. I would not do that. The only way you're getting me to lay down on a motorcycle is in a bikini for a calendar shoot.
Keep going for the video, head on the track happens about 20-seconds in if you have zero patience.
Thanks to my buddy c-nasty, who claims he can't ride a motorcycle without his penis dragging on the ground and getting run over by the car behind him.


text: In article <b8ad139e0904152318p5473cbe5yb5f55a19894cc834 at mail.gmail.com>, Jess Austin <jess.austin at gmail.com> wrote: > I'm new to python core development, and I've been advised to write to > python-dev concerning a feature/patch I've placed at > http://bugs.python.org/issue5434, with Rietveld at > http://codereview.appspot.com/25079. Without having looked at the code, I wonder whether you've looked at python-dateutil. I believe its relativedelta type does what you propose, plus much more, and it has the advantage of being widely used and tested. <http://labix.org/python-dateutil> -- Ned Deily, nad at acm.org


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text:
Hi all, if you have any questions or comments on the new B+ board, please add them here!
But first, my impressions...
The most noticeable addition is an extra two USB ports, which is achieved by replacing the old ethernet/USB 2 port hub chip with a four port version. Hand in hand with this is better power circuitry so the ports can now supply more power, which should help with hotplugging and high power devices (USB Hard drives should now work).
The SD card is now uSD, with a higher quality connector and less 'sticky outness'
All connectors are now on two sides of the board, and there are now mounting holes at each corner.
The GPIO connector now exposes more GPIO's (all of them I think!)
So, evolution rather than revolution, in order to maintain as much backwards compatibility as possible. Hopefully most of the points that people have complained about over the last couple of years have been addressed!


text:
Discover the Burgenlandkreis, located in the south of Saxony-Anhalt and be inspired by the spirit of the centuries. Amidst beautiful vineyards, you will find medieval castles and cathedrals. The "Naumburger Dom" is even UNESCO World Heritage.
The Burgenlandkreis is the most attractive district of Saxony-Anhalts. Castles, landscapes and nature reserves as well as the 60 km wine route and much more are waiting to be discovered.
Our house is quietly situated in the countryside and still the cities of Leipzig, Dresden, Weimar, Erfurt and Halle are in immediate vicinity. Spas, boat trips, lakes, cities of poets and philosophers, sports and socializing and (of course) finest wines are waiting for you.
You will find more information on our region at the Saale-Unstrut Tourism Board
The historic old city of Naumburg invites you to stroll through its narrow streets (guided tour possible) and will let you feel the spirit of centuries. Your way will inevitably lead you to the famous "Naumburger Dom", the Cathedral of St Peter & Paul, which is UNESCO World Heritage since 2018 and therefore one of the biggest attractions of the region.
Tradition and modernity are unified in the listed building of the largest sparkling wine producer in Germany – the Rotkäppchen Sparkling Wine Cellars in Freyburg. After an eventful tour you have an "exciting experience from thousands pearls" and taste a glass of Rotkäppchen sparkling wine. This fits especially our Rotkäppchen offer.
Visit Goethe's & Schiller's town of Weimar. Exemplary, start with Goethe's house with the National Museum. During a guided city tour, you will experience classic & contemporary and learn both, science and endearing about Weimar personalities: Goethe, Schiller, Bach, Liszt, Gropius and many more. After this interesting and insightful tour you have time to further explore Weimar on your own.
In contrary to the quiet town of Weimar there is the modern and lively city of Jena. Especially recommended is a visit to the Zeiss Planetarium. First opened in 1926, it is the oldest planetarium in the world. With a completely new, unique in this form, all-dome laser projection entertainment, knowledge transfer is offered in every season. A trip to Jena's Botanical Garden is always worth it.
Discover, one of the largest, nearly remained unscathed monuments from war. Explore the hometown and birthplace of George Friedrich Händel. At the market there is his memorial and at St. Mary's Church you can gaze the "queen of instruments", on which he once learned to play the organ music. The path also leads to the Cathedral, which was the first domain of the greatest son of the city.
In Halle another highlight is the oldest chocolate factory in Germany, the Halloren
Chocolate Factory. Experience the 200-year-old tradition of chocolate on a tour through the chocolate museum and the Chocolate Factory. The Halloren Chocolate Museum provides answers to many questions around the sweet seduction. Learn amazing things about the world of chocolate and the history of the oldest chocolate factory in Germany.
Visit to the Naumburg's cathedral, St. Peter & Paul
Sparkling wine tasting at the famous sparkling wine cellars of Rotkäppchen in Freyburg
Wine tasting sessions in the wine area Saale-Unstrut
Rallies to Leipzig, Dresden, Erfurt and Weimar
Knight's Dinner Neuenburg in Freyburg
Visit to the Horch Museum in Zwickau
Visit the Rudelsburg in Bad Kösen
Visit the playing cards Altenburg
Visit to the Käthe Kruse doll museum in Bad Kosen
Rafting on the river Saale from Naumburg
Dinghy or canoeing on the Saale or Unstrut from Naumburg
Ballooning from Osterfeld, Naumburg or Freyburg
Fahrsicherheitstraining at the Sachsenring in Zwickau
Kart race with award ceremony at the Sachsenring indoor kart track
Visit the Wasserburg in Schkölen
Visit to the Zeiss Planetarium in Jena
Visiting the Moritzburg in Zeitz
Visit the famous pottery Bürgel
Boat tours of a special kind with the Saale-Unstrut shipping company
This should be just a few examples of cultural and recreational opportunities in our region. Of course, we can also help you in planning your own personal ideas.
Event tips in and around the region of Naumburg you find at:
Our recommendation:


text: The EUR/USD pair initially fell during the course of the session on Friday, turning back around to form a bit of a hammer. The hammer of course is a very bullish candle, so if we can break above the top of a believe that the market should continue to go higher, perhaps reaching to the 1.13 level given enough time. Ultimately, this is a market that certainly looks like it's ready to break out to the upside, perhaps even going higher than we've recently seen and reaching towards the 1.15 level. We do not have any interest in selling at the moment.


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Whether it's a renovation, quality home, hospitality or light commercial project our company prides itself in working as a team with our clients to create a project that is tailored to meet their needs, functions well and looks great.
Our goal is to ensure the end result reflects the client's style rather than current trends.
We hope you enjoy this tour of some of our award wining homes and projects.
Copyright Mark Hawkins Architectural Design 2009. All rights reservedSite design by Media Suite


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Ballerina with bow, covered with glitter glitter to match both the jeans to trousers, ensuring a practical choice but tasty for everyday.
Bamboo socks with cute little fish; Super soft, absorbent (60% more than cotton), transpiring and antibacterial. Effective temperature control, leaving your feet cool in summer and warm in winter.
Comfortable suede uppers of shoe Derby, microfiber with dovetail joints. Lightweight model. Suitable for everyday: to give a touch of style to a sober and formal or to make it more elegant casual.
Thin wallet with coin pocket with zipper. Features 8 card slots inside, a large storage compartment and two slit pockets.
Vegan boots comfortable, flexible side zip closure. A simple and elegant design with soft Microsuede material. The elastic and side zipper makes donning easy. Top quality material and the non-slip resistant outsole provides maximum comfort, while the soft insole stitched in micronappa and the extra padding on the heel makes this model comfortable all day. With heel height 2, 5 cm.
Belt handmade in a small family-run factory in Portugal. With simple solid metal buckle (nickel free). Italian made of microfibre.
Vegan Sandals for women, Italian microfibre suede effect. Adjustable buckle strap with elastic for a perfect fit.
Beautiful soft Microfiber Jonaa Trapper hat with faux fur large band to wrap around the face. Delicious, warm and very comfortable, for a stylish colder periods.
Minimal jewelry holder, with a zipper around the appliance, at the top. Extra inside lining to protect your content.
Door-glasses structured with zipper closure and soft lining inside.
Décolleté ecological microfibre suede effect, with heel height 8 cm. Breathable and anti-allergic. Model always fashionable, versatile, suitable for different outfits.
Convenient two-tone suede upper, with Derby shoe of microfiber with dovetail, and insert in polyester with tweed. Lightweight model. Suitable for everyday: to give a touch of style to a sober and formal or to make it more elegant casual.


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New Year's Day 2014 has just arrived.
I'm not a big "make a resolution list" person. I think it's better to evaluate oneself all through the year.
But what I do like to do is think about what was good about last year, and dream about the New Year ahead. What do I hope to accomplish? Where will I go? What might be my happy achievements?
With my donors, I'm more focused. What do I hope to accomplish overall this year? What are my goals? Where will my donor(s) and I go on the philanthropic path? What do my donors hope to achieve and how might I help them get there?
This year I made a resolution. Tom Ahern handed it to me in his post, "What I learned in 2013."
"Deliver more gratitude than expected."
What is gratitude? Most people use this definition: "A feeling of thankfulness or appreciation." But I like this better: "Gratitude is paying attention to what we have, rather than what we don't."
This is doable. It is easy and yet a challenge. It may be applied to my fundraising and to my work with colleagues. And it works for my personal life too.
If you know me or have read my blog much, you know I believe in gratitude.
This is the perfect resolution for me: Deliver more gratitude than expected. How can you deliver more too?
For donors: Start with appreciation. End with thanks. And liberally sprinkle gratitude in between.
One place to start might be your thank you letters. Check out this link and this and this for tips to improve your thank you letters. And remember it is NOT a process. But it is one of the very early steps on the donor journey – one of the first steps you take together with your donor – so make it memorable.
How about a Thank-A-Thon? Or dip into last November's Nonprofit Blog Carnival, it was all about Giving Thanks and Gratitude. Or this Pinterest page…
Don't feel that everything has to be overhauled. Take one thing around thanks and improve it for your donors. Then another and another. And why not choose to thank a specific donor every week, above and beyond what you usually do?
For your team: Consider replacing frustration with gratitude. And give more than expected.
Feeling frustrated that you can't get Marketing team members "on board" with your donor-centered communications needs? Gnashing your teeth because Finance didn't handle a gift according to the donor intent? Longing for 100% board giving?
Instead of a hostile reaction (or worse, silently seething), professionally work through the differences in search of a solution. Then be on the look out for opportunities to thank your colleagues for a job well done.
Why? Happiness in the workplace increases productivity – and one way to do that is by recognizing achievements of coworkers. It can be as simple as sending a thanks for help to a colleague (even if it is expected) or writing a note of appreciation to leave on the desk – it can be as easy as your pen, a post-it and authentic words of gratitude. Or use an email, a fist-bump, a hat-tip in a meeting or a cup of coffee. There are lots of benefits of the Grateful Workplace.
For your personal life: Lead with gratitude once a day. Then try for more than once.
If you are in a relationship with a partner, you know (or recall) that gratitude plays an integral part of a healthy relationship. John Gottman has proven that couples with a high ratio of positive to negative interactions (5:1 or more) have the highest rates of intact, satisfying marriages (and this goes for committed relationships too!).
There are even benefits – physical, emotional and interpersonal – to keeping a gratitude journal. And if you have young ones in your life, you can teach them the art of gratitude as well.
My plan: I do have a personal goal to go with my resolution: to send a note, an email, make a call, give a hug every day before I sleep – all in the name of gratitude.
I'll be making a list of people to thank and seeking on-the-fly opportunities to express my appreciation, and to deliver more gratitude than expected.
What will you be doing in the New Year?
Yes! I want joy sailing through my inbox!
Open a package of smarts and joy! Every fortnight, find strategies, tips + inspiration to:
- Accomplish anything with grit, grace + gratitude.
- Navigate a successful career + balance your personal life.
- Create deeper relationships between supporters + your nonprofit.
- Understand how to work with high net-worth supporters.


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While many holiday goers in Europe seek the sun, sea and sand of the Mediterranean, it's scenery of a different nature that is order of the day during the winter months – ski slopes. From the Alps dominated skylines of France, Switzerland, Austria and Italy to the emerging ski holiday destinations of Bulgaria, Latvia, Romania and Slovakia, Europe has plenty of choice when it comes to ski holidays. Whether you're looking for a budget holiday, a once in a lifetime family break or a luxury chalet in the mountains, there's something to fit all budgets in the European ski scene. To help you get started when planning your next European ski adventure we've listed 5 of our favourite ski resorts in Europe.
5 Top ski resorts in Europe
1. Avoriaz, France
The ski resort of Avoriaz in the north of France may not be among the prettiest in Europe but it is full of character and more importantly, affordable. A popular family ski resort, Avoriaz has entertainment in the resort to suit all ages including Disney characters. The entertainment is great but you want to ski, and ski you shall with skiers and snowboarders falling in love with these slopes which aren't as challenging as others that France has to throw at you. A motor-free zone the only transport around Avoriaz is horse-drawn carts, just another part of the great character of this area.
2. Nendaz, Switzerland
The Nendaz ski resort is in the popular Valais region of Switzerland and offers skiers affordable access to the Four Valleys ski area, home of the epic free-ride terrain Verbier. With modern chalets and apartments available, you'll get to enjoy the off-piste
Nendaz in the Valais region of Switzerland offers back-door budget access to the Four Valleys, home to the epic freeride terrain above Verbier. Book into one of the modern but traditional and plentiful apartments in Nendaz and you'll be playing in the off-piste and itineraries above Tortin without having to pay through the nose for a Verbier chalet.
3. Baqueira Beret, Spain
When we think of a Spanish holiday we often think of the sandy beaches and sunny ourselves along the Costa's but the Spanish Pyrenees offer some wonderful ski slopes for skiers of all abilities. One of the most popular resorts in Spain is Salana in Baqueira Beret. With 115km of pistes and regular snow fall this area is particularly popular with stronger, more experienced skiers. With popular restaurants and some great accommodation, Baqueria Beret will give a Spanish holiday with a difference.
4. Soll, SkiWelt, Austria
If you're looking for a ski resort with plenty of choice and variety, both in terms of the slopes and things to do, look no further than the Soll ski resort in Austria. With over 250km of pistes to suit all abilities and 70 mountain restaurants, and nine ski villages, you'll never be stuck for things to do in Soll. Cheap beer and food in a perfect setting is an added bonus to the Soll ski resort.
5. Cervinia, Italy
The Cervinia ski resort in Italy is perfect for first time skiers and those of you with young kids looking to introduce them to a skiing adventure. It is often said that nothing compares to a ski holiday and if you are planning your first family ski vacation, you could do a lot worse than heading to Cervinia. This is a relatively small resort with lots of easy slopes for kids and more intermediate one's for the more experienced skiers among you. Cafes, restaurants and entertainment in the resort is perfect for families, making Cervinia a memorable family ski holiday.
If you're new to skiing holidays but want to experience them over the winter months then be sure to shop online to get the best deals. You can make some great savings by booking your own flights and accommodation through websites like Expedia but remember to book all of your skiing equipment in advance before arriving at your resort. This way you can check in and hit the slopes immediately. If you're a beginner be sure to avail of the training from the experienced instructors on the slopes.


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2016 Honda Odyssey Touring CARFAX One-Owner. Clean CARFAX. Local Trade, Sunroof/Moonroof, DVD, Leather Seats, Navigation.
This vehicle has been through a comprehensive Multi Point Safety Inspection that was performed by one of our Certified Mechanics. This vehicle will come with a 2 year maintenance program at no additional charge that will save you money! We also provide a 90 day / 3000 mile warranty that is specifically designed to give you even more peace of mind as you drive away in your new car - and this is in addition to the balance of the manufacturers warranty!
Call us today at 888-705-7938 or fill out the Get E-price form to get your very own special E-price! We can make this vehicle uniquely yours with window tint, custom tires, custom wheels, spoilers and DVD systems. This vehicle is available with our exclusive Car Doc Maintenance & Membership Program that is designed to give you peace-of-mind when it comes to taking care of your new car. We have the staff and knowledge to put it all together for you and make it your best car buying experience ever so CALL US NOW! Get Pre-Qualified: http://www.cookevillehonda.com/financing/application-short.htm We finance Bad Credit & First Time Buyers With No Credit. Apply Now! Or, Call Now (888) 705-7938.


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Comet Comics - Comic Convention - July 31st
Wed 29th Jun 2005
Tongue twisting stuf from Galwegian comic dealer and artist Emmet Taylor. There have been many rumours about 'planned' comics events this summer, and the first announcement is for July 31st in Wynns Hotel Dublin. This event will include a competition and for the kids a photoshoot with Spriderman, alsong with a slection of dealers. No guest announced as of yet.
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MindLab is a program for teaching problem solving and thinking skills with board games. My friend Avram, of Evolvems fame, turned me onto this program. I have not seen their program in action, but I met with the folks in charge of NYC MindLab, and was very impressed with their spiel.
The main idea is: you teach kids how to play any one of 200 games they have collected. Once the kids get the feel for the game and are struggling to master it, you step in and show them one of a dozen problem-solving methods MindLab has identified. They have cute names to help memorization and discussion. The kids learn the method quickly and then you move them to a second game. When the kids see this strategy works in two games, the method starts to stick in their brain. But the final and most important step is to transfer these methods to general life situations.
One method they described was the Stoplight method. The stoplight method is three steps: red, yellow and green. When you are playing your game, first do red: stop what you are thinking about (usually your next move) and think instead about our opponent. What is he thinking about? What move would he like to do. Yellow is to consider a counter-move. Finally with Green you choose your move, either a defense or an attack.
This method sounds fantastic—it's an vital strategy for any competitive game. But it can be applied so much more broadly. Imagine a conflict with a classmate. The "red light" step is to put yourself in your opponent's shoes. Consider how he sees the situation and figure out what his move might be. But this is exactly what conflict resolution courses teach—think about the other's position. Once you can see the other person's point of view, most conflicts can be resolved peacefully. Those that cannot, can be more successfully fought.
The stoplight method and dozens more are part of the MindLab curriculum. But in the end, it all comes down to the teachers. A great teacher will find teaching moments everywhere. The best curriculum in the hands of a poor teacher will just be boring with lots of wasted time.


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Rye Real Estate
The City of Rye is located in southern Westchester County on the shore of Long Island Sound, 25 miles from New York City, between Port Chester and Mamaroneck.
Approximately 15,000 residents enjoy 5.5 miles of water views, wildlife preserves, and open spaces. There is also a welcoming downtown area with shops, restaurants, nature center, and walking trails. It is separate from the town of Rye, which has more land area than the City of Rye's 6 square miles.
One-third of Rye's working residents commute to New York City; riding the Metro North Railroad from Rye to Grand Central takes about 45 minutes. Others are employed in Westchester, Connecticut, New Jersey, and Long Island as well as in the 200 small businesses and several large firms located here.
Despite its close proximity to New York City, the City of Rye retains its traditional atmosphere of tranquil village life, and Rye's remarkable natural endowments – a protected harbor along Long Island Sound, varied rolling landscapes, tree-lined streets, and winding brooks – enhance its many attractive neighborhoods.
FOCUS ON REAL ESTATE: Most real estate in Rye is made up of single-family homes that covers about three-fifths of the city's six square miles. Another fifth of the land is devoted to recreation and conservation. The balance is divided between institutions (such as The Osborn, a 56 acre retirement community, churches, and city property) and vacant land, with a slim 5% of all property in Rye used for business and industry. A big plus for Rye is that you get more land for your money than in some of the neighboring towns, such as Larchmont and Bronxville where zoning laws allow for denser construction.
The real estate market in Rye includes everything from (a small number of) condos and apartments, to split-levels, ranches, colonials, Tudors, and beautiful estates, some right on the Long Island Sound.
If you are looking for a beautiful home in a charming small city, we can show you all the City of Rye has to offer!
Click here for info on Rye's local schools


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Good cashflow management should be one of the most important priorities for businesses of any size. As has been so brutally demonstrated in recent months, it is possible for perfectly solvent companies to become unsustainable if cashflow becomes a problem.
If your company becomes unable to meet its short-term financial obligations, such as paying staff or suppliers, you may find it virtually impossible to remain in business without securing some kind of borrowing.
Poor Cashflow
The problem with poor cashflow is compounded by the UK's late payment culture. Customers leave invoices unpaid for as long as possible to boost their own cashflow, which has a knock-on effect on businesses which need to pay their own bills.
Furthermore, if customers don't pay their invoices in a timely manner, the growth prospects for a business are constrained. In this situation you would likely find it difficult to seize short-term opportunities as they arise, or make the capital expenditure necessary to expand.
As can be seen, the management of your cashflow is vital for both the short-term stability and long-term growth of your company. Developing strategies to ease your cashflow is important, particularly if you want to realise your business's full growth potential.
Invoice finance is a useful way for businesses of any size to secure their short-term financial position, and ensure that they have the resources on hand to take advantage of unexpected opportunities.
Invoice Financing
Invoice financing allows businesses to raise cash against the value of invoices that they have issued. The invoice finance provider, known as the factor, will pay a proportion of the invoice, often within 24 hours, and can then take on responsibility for ensuring that it is settled by your customer.
When the customer has paid, the factor will then pay you the remainder of the invoice's face value, less any administration charge.
There are a number of significant benefits associated with invoice financing. In the first instance, it offers some certainty regarding invoice payment dates. Rather than waiting 30 or even 90 days for payment, you can usually get up to 90% of the value of the invoice paid by the factor within 24 hours.
Factoring is therefore useful in the event of an unexpected expense or if, for example, a supplier offers a short-term deal of which you would like to take advantage.
Invoice finance provides your business with higher levels of working capital, an increased ability to make accurate financial predictions, and the opportunity to react quickly to changes in market conditions.
Furthermore, the credit that you secure through invoice financing will be directly related to the strength of your business; as your order book grows, so too will your credit line. This helps to provide you with the capital that you need to expand efficiently, quickly and in a risk-managed manner.
Invoice Finance Providers
There are a number of invoice finance providers, many of which also offer other forms of asset based lending. You should think carefully about your choice of provider, as most factors will expect you to commit to their service for a twelve month period.
However, if you find a factor that understands your business and is willing to support your plans, invoice financing can be an effective means by which you can manage your cashflow and grow your business.


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Welcome to HealingWell. This is a great site. I look forward to getting to know you better. This is Kitt.
I am sure you will find the members kind and caring and very supportive.
Please take some time and read through the threads on the site. I know we have some on dealing with the holidays.
Every day members of HealingWell recover their Self-esteem, their self-worth, their dreams, their pride, their dignity and a meaningful life.


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6th Asia Pacific Eldercare Innovation Awards 2O18 – Finalist
Emmy Monash is extremely proud to have been awarded three finalist awards at the 6th Asia Pacific Eldercare Innovation Awards 2018.
The awards were presented in Singapore and recognize exceptional aged care organisations for their innovative and quality approaches to change the way seniors age, and for their contribution in shaping the future of the ageing landscape.
The awards are part of International Ageing Week in Singapore which is an industry initiative driven by Ageing Asia, and comprises of a week of events, activities and training that showcase the world's latest innovations and demonstrate leading solutions that drive improvement in the quality of care for the older adults in the region.
CEO, Tanya Abramzon said, "To be recognised on an international stage for our innovation in active ageing programs as well as building architecture and interior design is an amazing achievement for Emmy Monash. We are immensely proud of this achievement and that we were able to showcase our facility and programs to a global audience".
Pictured are Tanya Abramzon, CEO and Judy Fetter, General Manager Operations with trophies for the three awards.
Emmy Monash was awarded finalist awards in the following three categories:
• Best Active Ageing Program: Still Firing
Journeys through Sculptures and Clay A partnership initiated by Emmy Monash Aged Care with Duldig Studios and Bialik College, Still Firing is a creative sensory program focused on clay sculpture techniques and personal expression. Residents learned new artistic skills, fostered inter generational friendships with the Bialik students and developed an appreciation of shared values and community wellbeing.
• Best Silver Architecture: Emmy Monash Gandel House
It's no surprise that the architectural design of Gandel House has attracted international attention since it's completion. We have created a community hub that encourages socialisation across generations, stimulates inclusiveness and offers opportunities for meaningful engagement. The architectural design allows for an abundance of natural light via vertically integrated inviting courtyards and themed lifestyle rooms to enhance interest based activities.
• Best Silver Interior Design: Emmy Monash Gandel House
To challenge the notion that quality of life diminishes with age, Emmy Monash Gandel House strikes the right balance of aspirational lifestyle with clinical care options in a positive ageing facility where residents live a full life; socially, emotionally, spiritually and physically in a refined urban landscape.
Emmy Monash Aged Care has become a vibrant community hub, dedicated to sustaining and advancing our focus on aged care services and residential facilities. The culture of embracing innovation whilst demonstrating respect, empathy, and excellence reflects our Jewish heritage and values. Encouraging volunteers, family, students, professional partners and staff to become actively involved with residents, reflects our holistic philosophy. Visionary leadership, committed board members, dedicated management, motivated staff and excellent facilities ensure residents needs are our paramount priority.
Innovative lifestyle programs enhance the imagination of our residents promoting self worth and quality of life, whilst the ageing in place structure encourages confidence, security and dignity for residents and their families.
Categories: News