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Extensive variation in synonymous substitution rates in mitochondrial genes of seed plants
© Mower et al; licensee BioMed Central Ltd. 2007
Received: 15 April 2007
Accepted: 09 August 2007
Published: 09 August 2007
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© Mower et al; licensee BioMed Central Ltd. 2007
Received: 15 April 2007
Accepted: 09 August 2007
Published: 09 August 2007
It has long been known that rates of synonymous substitutions are unusually low in mitochondrial genes of flowering and other land plants. Although two dramatic exceptions to this pattern have recently been reported, it is unclear how often major increases in substitution rates occur during plant mitochondrial evolution and what the overall magnitude of substitution rate variation is across plants.
A broad survey was undertaken to evaluate synonymous substitution rates in mitochondrial genes of angiosperms and gymnosperms. Although most taxa conform to the generality that plant mitochondrial sequences evolve slowly, additional cases of highly accelerated rates were found. We explore in detail one of these new cases, within the genus Silene. A roughly 100-fold increase in synonymous substitution rate is estimated to have taken place within the last 5 million years and involves only one of ten species of Silene sampled in this study. Examples of unusually slow sequence evolution were also identified. Comparison of the fastest and slowest lineages shows that synonymous substitution rates vary by four orders of magnitude across seed plants. In other words, some plant mitochondrial lineages accumulate more synonymous change in 10,000 years than do others in 100 million years. Several perplexing cases of gene-to-gene variation in sequence divergence within a plant were uncovered. Some of these probably reflect interesting biological phenomena, such as horizontal gene transfer, mitochondrial-to-nucleus transfer, and intragenomic variation in mitochondrial substitution rates, whereas others are likely the result of various kinds of errors.
The extremes of synonymous substitution rates measured here constitute by far the largest known range of rate variation for any group of organisms. These results highlight the utility of examining absolute substitution rates in a phylogenetic context rather than by traditional pairwise methods. Why substitution rates are generally so low in plant mitochondrial genomes yet occasionally increase dramatically remains mysterious.
A synonymous site substitution is defined as a change in a protein-coding gene that does not alter the amino acid sequence encoded by the gene. Thus, synonymous substitutions are often assumed to be free of selection at the protein level, and the rates at which they accumulate are widely used as an approximation of the neutral mutation rate. The first study on rates of synonymous substitutions in flowering plants found that mitochondrial genes evolve a few times more slowly than chloroplast genes, about ten times more slowly than plant and mammal nuclear genes, and 50–100 times more slowly than mammalian mitochondrial genes [1]. Later studies have confirmed the low rate of synonymous changes in angiosperm mitochondrial genes [2–4] and extended the observation to the entire mitochondrial genome [5]. However, recent studies identified two genera of flowering plants, Plantago and Pelargonium, that have experienced a dramatic increase in the mitochondrial rate of synonymous substitution [6–8]. Some of these rate increases were temporary, with rates approaching or returning to normally low levels in certain descendent lineages [6, 7].
Phylogenetic analyses have suggested additional cases of rate acceleration for several plant lineages [9–14], but it is unclear whether this is a widespread phenomenon in plants. Here we test the generality of slow synonymous sequence evolution in plant mitochondrial genes across a large number (between 306 and 578 species, depending on the gene) and wide diversity of seed plants (i.e., gymnosperms and angiosperms). Although genes from most species evolve slowly, as expected, additional cases of highly accelerated rates were found, as were examples of exceptionally slow sequence evolution. Surprisingly, a few plants were also identified that contain a mixture of both quickly and slowly evolving mitochondrial genes. Overall, these results demonstrate that the synonymous substitution rate in plant mitochondria is a more variable character than previously appreciated.
A surprising feature of these trees is the amount of branch-length variation seen (Figs. 1, 2, 3). Although the great majority of plants show conventionally low levels of synonymous site divergence, a number of species have substantially shorter or longer branches for one or more genes. The most striking examples of increased sequence divergence are from Plantago and Pelargonium, whose atp1 and cox1 branch lengths are exceptionally long (note that the matR gene has not been isolated from Pelargonium or from the fastest members of Plantago), as previously reported [6, 7]. Although less extreme, one or more species from Silene, Apodanthes, Acorus, Ephedra, and Podocarpus are also very divergent across multiple genes. Patterns are not as clear for other taxa. For instance, Goodenia and Musa are highly divergent for atp1 but only moderately divergent for cox1. For a number of species, such as Carex and Anthericum, a sequence from one gene is very divergent but data are not available for the other two genes.
In contrast to the examples of increased levels of mitochondrial sequence divergence, there are several groups that consistently show an unusually low level of divergence, even in the context of the general conservation expected for plants (Figs. 1, 2, 3). Within the angiosperms, these groups include the Arecales (also see [16]), the Chloranthales, the sister orders Laurales and Magnoliales, and most orders within the basal eudicots. Several gymnosperm lineages also have short branches, including the Cycadales, the Pinaceae, and Ginkgo.
Unexpectedly, these analyses have also identified several lineages with high levels of divergence for some genes but low levels for others (Figs. 1, 2, 3). For example, divergence levels for Alisma are high for atp1 and matR but unexceptional for cox1. The cox1 and matR genes from Ranunculus are remarkable in that they are unusually conserved, similar to the very slow rates found in most basal eudicots, while its atp1 gene is moderately divergent. For Polemonium and Pentamerista (in the Tetrameristaceae), atp1 is divergent but matR is conserved. Conversely, matR is divergent but atp1 is conserved for Pilostyles (in the Apodanthaceae).
The length of a branch in a phylogenetic tree, which represents an estimate of the amount of sequence divergence for that lineage, equals the product of the absolute substitution rate(s) and time. Because of this confluence, the rate and time components must be separated in order to make direct comparisons of substitution rates among taxa. We previously described a method to calculate absolute rates of synonymous (RS) and nonsynonymous (RN) substitution along each branch in a phylogenetic context [6, 7], and we employ essentially the same strategy here. However, in this case dS and dN values were estimated from a combined analysis of five mitochondrial protein genes (Fig. 4).
Divergence times, absolute substitution rates, and synonymous to nonsynonymous rate ratios for all terminal branches
Time (MYA)
P. lanceolata
17 ± 1
7.71 ± 2.08
0.53 ± 0.13
P. rugelii
17 ± 1
69.62 ± 6.33
2.76 ± 0.30
82 ± 4
0.19 ± 0.06
0.06 ± 0.02
89 ± 5
0.38 ± 0.06
0.17 ± 0.03
89 ± 5
0.10 ± 0.04
0.05 ± 0.01
S. latifolia
5 ± 1
0.54 ± 0.38
0.22 ± 0.12
S. noctiflora
5 ± 1
90.12 ± 13.94
5.37 ± 1.00
S. acaulis
8 ± 1
1.55 ± 0.52
0.45 ± 0.15
19 ± 4
0.59 ± 0.24
0.07 ± 0.06
38 ± 2
0.56 ± 0.12
0.04 ± 0.02
76 ± 4
0.29 ± 0.07
0.05 ± 0.02
41 ± 3
0.79 ± 0.72
0.06 ± 0.03
41 ± 3
30.70 ± 3.07
1.22 ± 0.17
100 ± 4
0.56 ± 0.07
0.16 ± 0.02
100 ± 4
0.70 ± 0.08
0.14 ± 0.03
100 ± 4
0.45 ± 0.07
0.09 ± 0.02
130 ± 6
0.08 ± 0.03
0.05 ± 0.01
131 ± 6
0.23 ± 0.05
0.03 ± 0.01
133 ± 6
0.11 ± 0.03
0.03 ± 0.01
133 ± 6
0.09 ± 0.03
0.02 ± 0.01
137 ± 6
0.36 ± 0.05
0.05 ± 0.01
86 ± 4
0.33 ± 0.08
0.03 ± 0.01
86 ± 4
0.61 ± 0.08
0.15 ± 0.03
92 ± 5
0.11 ± 0.04
0.05 ± 0.02
118 ± 6
0.19 ± 0.05
0.04 ± 0.01
127 ± 6
2.45 ± 0.21
0.26 ± 0.03
153 ± 8
0.26 ± 0.05
0.08 ± 0.02
158 ± 8
0.18 ± 0.04
0.07 ± 0.01
245 ± 0
0.43 ± 0.07
0.09 ± 0.01
245 ± 0
0.93 ± 0.10
0.08 ± 0.02
267 ± 8
0.47 ± 0.06
0.20 ± 0.02
286 ± 13
0.04 ± 0.02
0.13 ± 0.02
169 ± 27
0.38 ± 0.10
0.11 ± 0.03
169 ± 27
2.36 ± 0.43
0.31 ± 0.06
315 ± 5
0.10 ± 0.02
0.04 ± 0.01
179 ± 9
0.02 ± 0.01
0.03 ± 0.01
179 ± 9
0.06 ± 0.02
0.03 ± 0.01
It must be noted here that sites of RNA editing were not excluded in the analyses of this study. In plants, RNA editing usually occurs at multiple positions for any mitochondrial transcript [17, 18]. The retention of edited sites in rate analyses can sometimes bias estimates of nonsynonymous substitution rates, although their presence seems to have little to no effect on synonymous rate estimates [19, 20]. Indeed, the exclusion of edited sites in gymnosperm cox1 sequences resulted in a 30-fold reduction in the nonsynonymous rate [19]. Thus, the high RN/RS ratios observed for Pinus and Cycas are probably an artifact of the large number of edited sites present in their genes.
Number of RNA editing sites predicted by PREP-Mt (C = 0.5)
S. noctiflora
S. acaulis
S. latifolia
Previous studies on synonymous substitution rates found that plant mitochondrial sequences evolve very slowly [1–5]. For the most part, the results presented in Figures 1, 2, 3 confirm these earlier findings. Roughly 80–90% of the sampled species have normally low levels of sequence divergence at synonymous sites. However, a surprising number of species depart from the "normal" mitochondrial pattern. At one end of the spectrum, there are several angiosperm (e.g., Arecales, Chloranthales, and Laurales) and gymnosperm (e.g., Cycadales and Pinaceae) groups whose genes are unusually conserved at synonymous sites. At the opposite end are taxa such as Plantago, Pelargonium, and S. noctiflora with long branches for multiple genes. Many additional species have long branches for a single gene, but independent sequences are needed to confirm that these long branches are replicable and not artifactual (also see next section). On the whole, these results suggest that while most lineages of plant mitochondria do indeed possess generally low synonymous substitution rates, it's increasingly naive and misleading to categorize all plants in this way, and that synonymous substitution rates are, for reasons still unclear, exceptionally fluid and variable in plant mitochondria.
Because the analyses in Figures 1, 2, 3 used almost all species available in GenBank (some species were lost during data filtration; see methods), they provide a generally unbiased representation of rate variation across seed plants. However, for comparisons among species, branch lengths alone are inadequate because they are a function of absolute substitution rate(s) and time. In other words, equally long branches do not necessarily equate to equally fast absolute substitution rates. For example, dS for S. noctiflora, Acorus, and Ephedra are quite similar, but RS for S. noctiflora is almost 40 times higher than for Acorus and Ephedra (Table 1). Indeed, the synonymous rate for S. noctiflora is even faster than for the species of Plantago and Pelargonium examined here (Table 1), despite the much shorter branch length for S. noctiflora (Fig. 4).
These findings highlight the importance of taxon sampling in these analyses. The increased sampling of Silene species allowed us to pinpoint the rate acceleration in S. noctiflora to a 5 million year period. Similarly, sampling of multiple taxa within Plantago in a previous study on synonymous rates identified one species, P. media, with RS a few times higher than for S. noctiflora over a similar time span [6]. The finding of extensive variation in synonymous substitution rates within these two genera suggests that there is likely to be much more rate variation in plant mitochondrial genomes than is already apparent. Increasingly dense sampling of species is likely to uncover additional examples of rate variation. Calculation of absolute substitution rates may identify further examples, because some increases may be so recent that sequence divergence is still rather low and not readily apparent in phylogenetic trees.
Unfortunately, for Acorus and Ephedra there is little chance that increased sampling will provide a better estimate of the timing and magnitude of the rate acceleration, despite the fact that their increased divergence levels are spread over more than 100 million years. For Acorus, the genus encompasses only three or four species and they all appear to be very similar at the molecular level (see three species in Figure 1 and two species in Figure 3). Furthermore, there are no other genera in the Acorales to break up the time period from the base of monocots to the diversification of Acorus [22]. With over 60 species in Ephedra, it may be possible to more finely dissect the dynamics of rate evolution in this genus. However, like the situation in Acorus, the species included here show very little molecular divergence (see three species in Figure 2) and there are no other genera to break up the long time interval between the crown and stem group nodes of Ephedra [22].
Among-species comparisons reveal that synonymous substitution rates vary by four orders of magnitude across seed plants, from a low of 0.015 SSB in Cycas (Table 1) to a high of 166–244 SSB in Plantago media [6]. Excluding RNA viruses [23], this is, to our knowledge, by far the largest range of synonymous rates known for any phylogenetic group and genome, as well as both the lowest and highest estimated rates. To put this range in perspective, the amount of mitochondrial sequence divergence found in Cycas after almost 180 million years of evolution would have taken P. media on average only about 15 thousand years to accumulate. These results highlight the advantage of estimating rates in a phylogenetic context. Traditional pairwise methods average rates between two lineages, thereby masking any lineage-specific rate differences. A pairwise comparison between S. noctiflora at 90 SSB and S. latifolia at 0.54 SSB would result in an averaging of their very different rates to ~45 SSB, which doesn't reflect either of their actual rates very well. Similarly, the low rate found in Cycas (0.015 SSB) wouldn't be quite as low in a pairwise comparison with Zamia (0.059 SSB) leading to a pairwise rate of 0.037 SSB. The phylogenetic estimates of substitution rates indicate a 6000-fold range of variation between Cycas and S. noctiflora versus a 1000-fold range based on the pairwise estimates.
Although substitution rates are high for mitochondrial genes of S. noctiflora and Acorus, chloroplast and nuclear genes from these species do not show any increase in sequence divergence [see Supp. Fig. 1 in Additional file 1] [24–27]. Thus, the rate acceleration in these species is confined to the mitochondrial genome, as seen also in Plantago and Pelargonium [6, 7]. In those studies, defects in mitochondrial DNA replication or repair were speculated to be, among a wide range of processes potentially affecting the mitochondrial mutation rate, the most likely causes of such severe, mitochondrial-specific increases in substitution rates. Similar factors may also be at work in S. noctiflora and Acorus. In contrast, in Ephedra, chloroplast and nuclear sequences are also divergent [27, 28]. This points to a factor acting at the organismal level, such as generation time, that would affect rates in all three genomes, although mitochondrial-specific forces may be at work as well. Some studies [16, 29–31] have detected moderate, correlated differences in synonymous substitution rates across all three plant genomes and have attributed these to generation time effects [16, 29, 30], paternal transmission of organelles [31], or correlated substitution and speciation rates [32, 33].
One surprising finding from the broad-scale analyses is the number of plant lineages with levels of synonymous site divergence that appear to be relatively high for some genes and low for others (Figs. 1, 2, 3). In most cases, the basis for this apparent within-plant rate heterogeneity is unknown, and as emphasized below, some of these cases are likely to be or potentially are the result of error of various kinds. Others, however, have a possible biological underpinning. One likely biological explanation is horizontal gene transfer, which occurs surprisingly frequently between mitochondrial genomes of unrelated plants [34, 35]. Transfers between genomes with highly different substitution rates and accumulated sequence divergence could easily account for some of the within-plant, gene-to-gene differences shown in Figs. 1, 2, 3 and indeed is suspected to account for one particular case. Two members of the Apodanthaceae, Pilostyles and Apodanthes, have high levels of divergence for matR. The Apodanthes atp1 gene is similarly divergent, but atp1 from Pilostyles is not. Phylogenetic analysis of the atp1 gene from Pilostyles suggests the possibility that this gene may have been acquired via horizontal transfer from a plant with normally low mitochondrial rates [12].
Another potential biological explanation for differences in synonymous rates between genes from the same plant is that they are located in different genetic compartments that possess different substitution rates. With the exception of Plantago and Pelargonium [6, 7] and probably some of the new cases of high-rate mitochondrial genomes reported in this study, synonymous rates are much lower in mitochondrial than nuclear genomes in plants [1–4]. Functional transfer of a mitochondrial gene to the nucleus will thus usually lead to much higher rates of sequence divergence. This possibility is exciting because there has been no report of functional nuclear transfer for atp1, matR, or cox1 in any plant. This includes no evidence for loss of these three genes from the mitochondrial genomes of 280 diverse angiosperms examined by Southern blot hybridization, whereas 16 other genes were inferred to be frequently lost from the mitochondrial genome and, equally frequently, functionally transferred to the nucleus [36].
A third intriguing possibility is that synonymous substitution rates vary across regions of the mitochondrial genome. In the chloroplast genome of angiosperms, synonymous rates are known to be a few-fold higher in single-copy regions than in a large inverted-repeat region [1, 37], and of perhaps greater relevance, a small region with higher accelerated synonymous rates has recently been discovered in the chloroplast of one lineage of legumes (KH Wolfe, personal communication). Gene-to-gene variation in the mitochondrial mutation rate was also recently observed in populations of Silene vulgaris [38, 39].
There are also several non-biological reasons why divergence levels for a particular species vary between genes. Sequencing errors will artificially inflate sequence divergence. The Pennisetum atp1 sequence is a likely example. Including the atp1 sequence, eight mitochondrial genes were generated for Pennisetum from the same unpublished study (GenBank: AF511559–AF511569), and all show similar characteristics including frameshifting indels and a large number of nucleotide substitutions (data not shown). Most likely, these unpublished sequences are full of errors, and this is the cause of the apparently anomalous divergence of atp1 in Pennisetum.
Misidentification of the correct phylogenetic position of a sequence/organism could also lead to artifactually long branches (recall that topological constraints were enforced for dS branch length estimation). For example, two cases were found where the taxonomic sources of sequences were swapped. In the first case, a cox1 sequence whose source is listed as Austrobaileya (GenBank: AF193954) is more similar to an independent Ceratophyllum sequence than to other Austrobaileyales sequences, while a sequence annotated as Ceratophyllum (GenBank: AF193945) is actually more Austrobaileya-like. These two GenBank files have now been corrected. A similar mix-up occurred between a Ginkgo-like sequence annotated as Cabomba (GenBank: X94585) and a Cabomba-like sequence identified as Ginkgo (GenBank: X94587). In each example, the mislabeled sequences were generated from the same study and probably reflect a mix-up that occurred during the database submission process. They were easily identifiable (and were excluded from the final analyses) because they appeared as uncharacteristically long branches relative to independent sequences of the same gene from the same genus.
The use of topological constraints in the analyses may similarly lead to spuriously long branches. To evaluate this possibility, unconstrained analyses were performed on the four data sets used for Figures 1, 2, 3, 4 [see Supp. Figs. 2-3 in Additional file 1]. As can be seen, almost all long branches in the constrained analyses remain long in the unconstrained analyses. However, the use of topological constraints apparently led to an erroneously long branch for Archytaea matR and for Pennisetum atp1. Archytaea was constrained as an asterid in the Ericales (based on GenBank taxonomy), but it is actually a rosid in the Malpighiales [22]. In the unconstrained analysis for matR, Archytaea groups properly within the Malpighiales and exhibits a more normal branch length [see Supp. Fig. 2 in Additional file 1]. The long branch for Pennisetum atp1 also greatly diminishes in the unconstrained analyses [see Supp. Fig. 2 in Additional file 1]. As mentioned above, the long Pennisetum branch may result from a poor-quality sequence read. Alternatively, the sequence may have been acquired via horizontal gene transfer, or the DNA sample was derived from an organism mistakenly identified as Pennisetum.
These examples underscore the need for independent sequencing to verify that any and all cases of putative intragenomic variation in synonymous substitution rates are real and not the result of human error. For those cases that are validated, it will be interesting to see whether any turn out to reflect noteworthy events in mitochondrial evolution, such as gene- or region-specific differences in substitution rates within a mitochondrial genome, horizontal gene transfer, or the functional transfer to the nucleus of a gene that has never known to have been so-transferred in plants (atp1 or matR) or even among all eukaryotes (cox1).
In this study, we have measured synonymous substitution rates in a phylogenetic context and uncovered numerous independent examples of rate increase and decrease. These results demonstrate that the synonymous substitution rate in plant mitochondria is a more variable character than previously appreciated. The extremes of synonymous substitution rates measured here constitute by far the largest known range of rate variation for any group of organisms, yet there is no obvious explanation for these divergent patterns. Future studies are required to understand the evolutionary processes driving these patterns.
Total genomic DNA was extracted and purified from fresh leaves using a CTAB protocol [40] or using the DNeasy Plant Mini Kit (QIAGEN) according to manufacturer's instructions. Genes were amplified by polymerase chain reaction using a PTC-200 thermocycler (MJ Research) and gene-specific primers [see Supp. Table 1 in Additional file 2]. Each reaction was performed using 35 cycles of 30 sec at 94°C, 30 sec at 50°C, and 2.0–2.5 min at 72°C, with an initial step of 3 min at 94°C and a final step of 10 min at 72°C. Polymerase chain reaction products were purified using ExoSAP-IT (United States Biochemical) and then sequenced on both strands using an ABI 3730 (Applied Biosystems) at the Indiana Molecular Biology Institute. Sequences newly determined in this study were deposited in GenBank under accessions EF547202–EF547251. Additional nucleotide sequences used in this study were obtained from GenBank [see Supp. Tables 2-7 in Additional file 2].
Similarity searches were performed with blastn against the nonredundant GenBank database using the atp1, cox1, and matR genes from Arabidopsis thaliana as queries (GenBank: Y08501). Limitations were enforced during the searches such that only hits from the Spermatophyta (gymnosperms and angiosperms) with an e-value less than 1e-10 were included. Hits less than 200 bp in length, multiple hits to the same species, pseudogenes, and cDNA sequences were removed from the blast results, while sequences generated in this study were added. Sequences were aligned using ClustalX and manually adjusted when necessary. Poorly alignable regions were excluded from the data sets, as were regions with gaps in the majority of taxa. These data sets (and all others used in the paper) are available [see Additional file 3].
For each surveyed gene, a ML topology was determined with PAUP*. To ensure completion of the ML analysis, several time-saving steps were taken. First, a partially-constrained topology was enforced. The topology was constrained such that all sequences within a family (as defined by GenBank taxonomy) were forced to be monophyletic. Relationships among families were constrained according to information from the Angiosperm Phylogeny Website [22]. Relationships among the major seed plant groups were constrained according to the 9-gene ML analysis of Qiu et al [41]. Second, ML parameters were fixed to values estimated from a neighbor-joining prior to the ML analysis. Third, the ML analysis used the HKY+G+I model with four rate categories rather than the more parameter-rich GTR+G+I model. The ML topology was identified by a heuristic search starting from the aforementioned neighbor-joining tree and using the TBR branch-swapping algorithm and the MULTREES option.
To determine whether the PAUP* time-saving strategies had an effect on the results of this study, a ML topology was also determined for each gene using RAxML version 2.2.3 [42]. In contrast to the PAUP* analysis, no topological constraints were enforced, the GTR+G model was used (GTR+G+I is not available in RAxML), the search started from a maximum parsimony tree, and the ML parameters were estimated during the ML analysis. For each gene, 10 replicates were performed starting from 10 different randomized parsimony trees. The initial rearrangement setting was fixed to 10 for all analyses.
Using the ML tree topologies generated above for each gene, dSbranch lengths were estimated in HyPhy version 0.99 for UNIX [43] using the MG94W9 codon model [15] and allowing for independent dN and dS values for each branch (the local parameters option). The dS trees using the PAUP* topologies are shown in Figures 1, 2, 3, and the dS trees based on RAxML topologies are shown in Supplementary Figure 2 [see Additional file 1].
Absolute substitution rates were calculated for a subset of the taxa in the rate survey using methods that have been described previously [6, 7]. Briefly, divergence times within Caryophyllaceae [see Supp. Fig. 1B in Additional file 1] were estimated by penalized likelihood with the program r8s [44], using an ML tree from an analysis of the chloroplast gene matK [see Supp. Fig. 1A in Additional file 1] and 38 million years for the split between Caryophyllaceae and Amaranthaceae as a calibration point [45]. Standard errors for Caryophyllaceae divergence times were estimated from 100 bootstrap replicates. Divergence times and associated errors for Plantago, for the remaining angiosperms, and for gymnosperms were taken from the analyses of Cho et al [6], Wikstrom et al [45], and Magallon and Sanderson [27], respectively. The dS and dN trees in Figures 4 and 5 were estimated from individual and combined data sets of five mitochondrial protein-coding genes (atp1, cob, cox1, cox2, cox3) using HyPhy as described in the previous section on a topology constrained according to the Angiosperm Phylogeny Website [22]. Standard errors for dS and dN were estimated from 100 bootstrap replicates. RS and RN values for each branch were calculated by dividing the mitochondrial sequence divergences from the combined data set by the elapsed time along that branch (Table 1).
The authors thank Chris Parkinson for providing the Alisma cox1 sequence and Sergei Kosakovsky Pond for helpful assistance with the HyPhy software. This work was supported by NIH research grant GM-70612 (to JDP), NSF grant DEB-0210971 (to LFD), and a Fulbright/Nord Pas de Calais Region Award (to PT).
This article is published under license to BioMed Central Ltd. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/2.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.


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MINNEAPOLIS -- Eight days after groin surgery, Minnesota Vikings running back Adrian Peterson is making the rounds in New York City before the Super Bowl. He's been conducting interviews for a handful of promotions, headlined a Super Bowl party earlier this week and is hosting a comedy show featuring Chris Tucker in New Jersey on Friday night. On Saturday, he'll present the Comeback Player of the Year award at the NFL Honors show.
Peterson stopped by ESPN Radio's "Mike and Mike" show on Friday morning, and said he's feeling fine after surgery, adding he's been able to do some upper-body workouts and will be back to normal in about six weeks.
"I would probably be back working (out already if not for the surgery, but) not too much. It's not going to interfere," Peterson said. "I'm still able to do some of the upper body exercises and things like that, but kind of laying off the lower body workouts."
The running back said again that he's keeping an eye on the Vikings' moves this offseason, and mentioned how gratifying it was for players to name him the player they'd most like to see in a Super Bowl. But as we've discussed, Peterson typically isn't one to hang onto his objections, at least publicly. On Friday, the running back -- who was upset the team fired coach Leslie Frazier on Dec. 30 -- gave the strongest endorsement he's given yet of new coach Mike Zimmer.
"I'm just looking forward to heading in the right direction. I feel like the organization did a great job of bringing in Zimmer and offensive coordinator Norv Turner," Peterson said. "I feel like we're putting the pieces in the right place so I'm just excited to see who all we bring in (to coach) other positions and to see which direction as far as bringing in players to help us accomplish our goal of winning a championship."


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he was just using a sleeping bag with not sure the rating of the bag.
Type: Posts; User: Can't Wait
he was just using a sleeping bag with not sure the rating of the bag.
Sold thanks for looking
PM sent sold pending funds
Just a little bump
Used maybe 4 times MSR Whisperlight international stove includes all tips accessories and 20 OZ fuel bottle $60 shipped lower 48 Paypal
Last price drop $30 shipped collecting dust thanks for looking
Bump would like to get these out of my way
Price drop great pair of gators at a fair price OR Verglas Gators size large Exc cond. $35 or shipped lower 48 Papal , sorry for yesterdays screw up on my part
Bump folks give me...
Please read update above Thankyou
Will accept reasonable offer on package deal to save on shipping
Have updated price as package let me see if any interest first save me money in shipping
Just a Bump
OR Short Gators Large Exc Cond $25
OR Verglas Gators Tall/LRG Exc Cond $40
eVent Mitts Never worn seam sealed $35
MSR Whisperlight International stove with repair kit and 20oz...
Sold please lock post
Sold pending funds PM for Paypal info Steelwolf
Paypal Excellent shape No Stains holes Tares. includes 20' Speer no tangle ridge line, 8' tie outs with prussic 4 ea for adjustment 4 ea side pullouts with bungee and 10' ea Speer no...
Sold please lock
No rips tears stains hung inside smoke free home Well cared for gear
Bump sorry for wrong model to many ULA packs
OOPs your right Catalyst bought a Ohm haven't used this one in 3 year so 4 years old will correct post
Large torso. I'm 6' 240lbs also has bladder pouch and gear pouch
For sale ULA Catalyst Great condition $150 shipped lower 48 Paypal
Large torso found the tag I was 21
Will post pics probably 2 years old used dozen times I want to say torso is in 20s I am 6'
Withdrawn no longer for sale
Sale withdrawn
Would have to pay for extra shipping costs
Yep still for sale bump
Just a little Bump
Sold thanks CST and everyone for looking
Bump included shipping lower 48 sorry for confusion
Sold CSt pending funds did know that fee was paid by seller
It's all Black no draft tubes older suspension
Hammockgear Black 0° full length incubator bought 2 years ago used 6 times hangs uncompressed in home. $200 shipped CUS Paypal buyer pays fees thank you
Brand new MSR axis snowshoes 30 " wore twice just to try no damage whats so ever have bracket for walking up and down hills adjustable gate and easy on and off. Just collecting dust sell $200 ...
I think I am just going to try to sale local it's just a royal pain to ship stuff of this size out and cost I and giving this stuff away at this price as it is don't want to pay more for shipping .
I appreciate that my friend but packaging and shipping a pulk is a royal pain in the arse lol
Wow no ones interested bump lol
Will keep you in mind
Withdrawn Closed
Well my best has been 2° F , I could probably do better , but after having a pack liners freeze where I couldn't put them on and just frozen gear in general I switched to hot tenting for deep cold....
All sold. Will be cleaning out more gear soon
Filter sold pending funds
Down booties sold. Water filter great shape have too much gear
I will pm you with paypal info soon as by pc booties sold pending funds
Goose Feet size large worn once $50 shipped
Katadyn Hiker Pro filter with extra new filter $ 50 shipped filter...
Ya I maid a loop out of armsteel to use as a extender see if that works
What I was hoping. I know wondering fool was getting hung by his as well Nice to get ti out of the way. Thanks my friend.
Hello been awhile guys and gals question. Bought another WBBB my old one got a hole in bottom patched it but leary on using way back in. Well I always hated trying to sit and recline in it with the...
thanks folks i am just trying to protect our sport


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"one of the most wonderful places in the world" Top 5 Page for this destination Lanzhou by gmanxue
Lanzhou Travel Guide: 99 reviews and 188 photos
A wonderful place I ever have been. Full of the legends of the Northwest in this place. In antient China, merchants went aboard to trade their products, so Gansu became an important place they must go through and became the main part of silkroad. Because of this history, there are lots of stories and culture reserved from ancient China. In Gansu Province the most famous landscape is Dunhuang Cave, in which are full of the murals. All the murals are more than 2,000 years old, and now you can still see the figures clearly. There are about 1,000 caves and innumerable murals there .This pic is about the white elephant. You can see its fresh color and vivid designings.
More than 5,000 years ago, ancient Chinese people began to live along the Huanghe River. As every Chinese, the Huanghe River is the cradle of Chinese civilization and the river of life. Because a great amount of sand depositting on the river bed, the color of the river is yellow. The Huanghe River goes across Lanzhou, so you may see the quiet part of the river and grand bridges. Everytime I see her, I always feel excited and proud.
Reviews (11)
The special nightlife in Dunhuang is eating outside. Because Dunhuang has a long winter, there always some small tents... more travel advice
Here is a list I think worthy to buy. Antique--- Goblet made by black jade, flying horse made by copper, some ancient... more travel advice
I am lucky enough to see how to make Ye Guang Bei. The best place to see it is Jiu Quan, Chinese Satellite Base. There... more travel advice
The ancient dance of Dunhuang revives from the Dunhuang freso. Professionals translate the dance on freso to our real... more travel advice
Comments (19)
Great page. Looking forward to visit Lanzhou in a few weeks.
Nice pages, nice phtographs and tips..Many many happy returns of the day.Wishing you a happy birthday...Have a nice time....
great page - gotta love those camels :-)
magnificent photos-very beutiful page
the mural are closer through VT but I must see them soon ;-))
great page
Superb pictures
Great page!
This looks a wonderful area especially those caves
Great comments and pictures
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Students who are currently enrolled in medical school and considering dual-degree training should apply to the NIH MD/PhD Partnership Training program and to one of the NIH Graduate Partnership Programs (GPPs), such as the NIH Oxford-Cambridge Scholars Program (OxCam).
As a Track 2 student, you are responsible for funding the pre-clinical years of medical school (usually MS1 and MS2) and will begin the PhD phase of your training in an NIH GPP. If you are accepted into one of the PhD programs at the NIH, you are conditionally awarded supplemental NIH MSTP funds, available through the NIH MD/PhD Partnership Training Program, to support your remaining years of medical school (usually MS3 and MS4). If you are interested in creating an Individual Partnership, you will need to contact the MD/PhD Partnership Training Program and the NIH GPP.
For students who are interested in funding for the remainder of medical school through the NIH MD/PhD Partnership Training Program, please answer the following questions for the steps in the application process:
NOTE: MSTP is defined here as a Medical Scientist Training Program with a T32 grant. A list of schools that receive the MSTP T32 grant can be found here: http://www.nigms.nih.gov/training/instpredoc/pages/PredocInst-MSTP.aspx
Students who are already enrolled in their medical school's MSTP must complete the following in order to participate in the NIH MD/PhD Partnership Training Program:
NOTE: Even if you are not awarded supplemental NIH MSTP funding, you are still encouraged to participate in activities and events planned for MD/PhD students at the NIH through your graduate program, the GPP, and the NIH Clinical Research Center.
Individuals who are enrolled in a medical school that has an MSTP, but are not enrolled in that program, must complete the following in order to participate in the NIH MD/PhD Partnership Training Program:
Students who are enrolled in medical schools that do NOT have an MSTP are, unfortunately, not eligible for supplemental NIH MSTP funding. Please feel free to apply to any of the options available through the NIH Graduate Partnerships Program (GPP). The activities and events organized for students by the GPP, individual programs, and the NIH Clinical Research Center are open and available for you. (You must speak with the Dean of your medical school about your intentions and get permission to participate in one of the NIH partnership programs for the PhD portion of your training.)
At this time, most students in the NIH MD/PhD Program earn their PhD in OxCam, a four-year PhD program partnered with the Universities of Oxford and Cambridge. For more information about OxCam, please visit its website.
If you are considering a different NIH GPP or Individual Partnership, please contact the program administrators or the main GPP office for more information.
Only the NIH GPP application is required for considering your admission to the NIH OxCam Program or other NIH GPPs:


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Neil Armfield's film hits hard because it sensitively shows how life on drugs can never be about anything else, and how the real horror of addiction is not what users do to themselves, but what they do to each other out of loneliness and despair.
For all its depiction of a descent into drug addiction, Candy is filled with surprisingly sweet moments and goes down more easily than seems possible given the subject matter.
With its intelligence, compassion, human terror and sheer loveliness, Candy is a winner despite the well-worn path it treads.
Doesn't add anything substantively new, though it has been nicely directed by Neil Armfield, known in his country for his theater work, and features striking performances from Heath Ledger and Geoffrey Rush.
Though it's a well-worn story, Candy does touch on a universal anxiety. For two people basking in the heat of an all-consuming love, what happens when the power gets cut off?
Because although there are some very striking moments in Neil Armfield's debut, there are simply not enough to keep us absorbed the way a movie should.
The way director and co-adapter Armfield shoots it, the film's awfully pretty in its grimness, in the way "Leaving Las Vegas" managed to make train-wreck alcoholism more fake-lyrical than grungy.
As the characters' lives fall apart, Ledger fails to bring the necessary gravitas to the role, and he looks a bit too healthy throughout.
Life, love and addiction make a mostly bitter, but occasionally sweet, concoction in Oz drama Candy, which is sometimes hard to swallow.
Cornish provides a counterbalance for Ledger's authoritative presence, turning what could have been just another heroin movie into a flawed but engrossing parable on love and sacrifice.
Ledger mumbles his entire performance (some of it barely legible) as a fuzzy, friendly, happily passive heroin addict and sometime poet, as if he's too blissed out to even open his mouth as he simply drifts along with his addiction.
Any drug movie's effectiveness can be measured by the strength of its detox, and Candy doesn't sweeten the cold turkey. Still, it's a downward spiral from there in more ways than one. Never mind the neo-psychedelic-pop soundtrack and occasional double-vision cinematography: Dope just can't account for the film's fried brain cells.
As the movie's feet get stuck in its own misery, it made me appreciate "Trainspotting" all over again - its wit, how it moved, the way any outcome for its characters seemed possible.
Ledger turns in another stellar performance and Cornish is heartbreakingly good also in this well-crafted film. But once that first plunger is pushed, the surprises are few.
Geoffrey Rush is fine as a gay drug dealer who serves as an enabling Santa Claus to the doomed couple. But in the end, Candy is a little too sweet and not quite harmful enough to the audience's health.
There has to be a good reason to put yourself through yet another junkie odyssey and Candy flunks the test.
What we see, though, is the same old same old - beautiful faces turning gaunt and haunted, strung-out hero and heroine, stupid parents, de-tox worse than tox, descent to and return from the depths. Candy could be seen, I suppose, as a cautionary tale; take this as a cautionary review.
Decent performances aside, the only interesting bits involve Geoffrey Rush as a chemistry professor who enables their self-abuse.
It's neither utterly real nor utterly romantic (heroin, like alcohol, manages to be awfully and unremittingly both).
User Score
Generally favorable reviews- based on 29 Ratings
User score distribution:
Positive: 4 out of 4
Mixed: 0 out of 4
Negative: 0 out of 4
Oct 10, 2015This film is brilliantly acted and is well supported by a riveting story. As the film goes on you become more attached to the characters, andThis film is brilliantly acted and is well supported by a riveting story. As the film goes on you become more attached to the characters, and will most likely be devastated yet filled with emotion at the end. As the credits rolled and the music played, I got goosebumps and contemplated how it must feel inside the body of an addict.… Full Review »
ChadS.Mar 31, 2007Being a painter, when Candy (Abbie Cornish) moves to the countryside with Dan(Heath Ledger, who reclaims his heterosexual screen image in a Being a painter, when Candy (Abbie Cornish) moves to the countryside with Dan(Heath Ledger, who reclaims his heterosexual screen image in a scene that makes ironic use of "My Own Private Idaho"), Jackson Pollock and Lee Krasner's migration to the boonies spring to mind. But then you realize that Dan doesn't do anything, so the prescient evocation becomes only half-right. Dan is no action painter; action hay bailer, yes. Candy can still be Krasner, but her influential(the bad kind) beau is the anti-muse. Dan has nothing to offer her, except his love. Is it enough? No. Candy may, or may not mature into an artist of great notoriety, but she'll never get a chance to find out if the heroin does her in. "Candy" is a love story about co-dependents(who might be star-crossed; she's middle class, he might be "white trash") that has its intermittent moments of underlining the debacle, which is the grade of diaster that drug addiction entails, with startling power and heartbreak. The sequence that documents Candy's attempt to bring life into this world stands out from the rest of "Candy" in which, like all movies about addicts, are broken into two sets of scenes; they're either stoned, or not stoned.… Full Review »
Helenp.Nov 30, 2006It's simply gorgeous!


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For Rent, island home on North Captiva, away from it all
Gulf Coast island located just north of Sanibel and Captiva Island.Caribbean like island 3½ miles long and ½ mile wide at the widest point.
4 miles of continuous natural beach.
Over half the island is state nature preserve (our house has wonderful views of the preserve).
Leave your cars on the main land. Electric golf carts available on the island for transportation. Use of one golf cart included in the rental. 2nd cart available for $250.00. Convenient water taxi service starts your adventure with a beautiful 20 minute boat ride.
Island has 3 restaurants, 2 gift shops, a pool bar and a small convenience store.
Legend has it Captiva Island was named by the pirates who frequented the Gulf Coast and held hostage beautiful women giving the island its name Captiva. North Captiva was part of Captiva Island until a hurricane in 1921 created Redfish Pass, separating the 2 islands.
This tropical paradise is filled with natural wonders and the island pace to life
The balance between man and nature is an island phenomenon. Birds, tortoises and other wildlife abound.
Some of the best shelling in the US is on our beaches. (Ask about the best shelling locations when booking the house.)
Good fishing from the beach or charter boat.
Captivating sunrises and sunsets.
Free kayacking in the back bay. Excellent for birdwatching especially at low tide.
All of this can be enjoyed from your own tropical veranda
New 4 bedroom/4 bathroom reverse floor plan pool home (click to see floor plans) just a short stroll down a sandy lane to the beach. Great views of State Preserve and Gulf of Mexico from its veranda and decks.
Verandas also provide space for outdoor dining and relaxing, watching the sunset over the Gulf in one of the incredibly comfortable adirondack chairs.
The ground level imported tumbled marble patio extends out to the heated pool providing space to gather on comfortable patio furniture and access the outdoor kitchen and dining table.
Gourmet Fully Stocked Kitchen with stone counters and
stainless steel appliances.The kitchen/living room veranda provides a wonderful space for dining and watching the sunset and sunrise.
Rain head showers in both master bathrooms. Whirlpool tub in kids bathroom.
Loft with sofa couch and sundeck.
Cathedral ceiling in the living room and dining room fill the space with light and openness.
Interior of home is professionally decorated with high end furniture and truely wonderful color scheme.
Bring your laptop and connect to our DSL wireless system. Check your email or surf the net while sitting by the pool or from the veranda overlooking the Gulf.
Guest Membership to Island Club included in rental.
Island Club membership entitles guests to use all Island Club facilities:
For Photo Gallery, click here or on one of the photos below or check out our video


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Citizen journalism image provided by Coordination Committee in Kafr Susa which has been authenticated based on its contents and other AP reporting, shows people tearing down a huge poster of President Bashar Assad and hitting it with their shoes, in Raqqa, Syria, Monday, March. 4, 2013. The activists said the picture was taken inside the Air Force Intelligence headquarters in Raqqa. (AP Photo/Coordination Committee In Kafr Susa)


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This is a Portable version of Mozilla Firefox with several add-ons that are useful for Web Application Security. The purpose of this package is to have the best available addons to manually test XSS, SQL, siXSS, CSRF, Trace XSS, RFI, LFI, etc.3 weekly downloads
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H323 Auto Caller is a simple H323-based WAV file player: you use it to call a H.323 telephone and it will play a WAV file to the remote telephone over the H.323 connection. Possible uses include Voice alerts with Nagios/OpenNMS and hotel wakeup systems1 weekly downloads
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Like the well-known tee program on UNIX system, httpTee is sitting between the client and the server of a HTTP / HTTPS connection. Allows the requests and responses to be modified before submission. Great for web application development or students.
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Dictionary and affix files for the aspell, myspell, hunspell and ispell programs for Ukrainian language. Словники для перевірки правопису української мови за допомогою aspell, myspell, hunspell16 weekly downloads
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Hylafax Job Control Langauge (HJCL) is a batch processing perl script front end for Hylafax using PERL MAGICK and IMAGE MAGICK. Scans a directory for fax jobs and submits them to Hylafax.1 weekly downloads
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ispCP is a project founded to build a Multi Server Control and Administration Panel without limits. This Control Panel should be usable by any Internet Service Provider (ISP). This project is a fork of the stalled VHCS.65 weekly downloads
Indytrans integration server
Integration server integrates critical business systems. Connects to database supporting OleDB, ODBC og native MS Sql server, XML, textfiles, custom sources with C# scripting. The solution includes a library, Windows service, and a configuration client.
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Various shellscripts to automate process of installation and chrooting of both NaviServer and AOLserver Webservers
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A distributed database of XML documents for Intranets and the Internet in a region where connectivity is sporadic. Development is by UN Volunteers and Loja Technical University, to support staff of the Business Development Agency in Loja, Ecuador1 weekly downloads
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Extracts information about jsp pages for use by developers. Integrates with regular javadoc documentation for .java files.
JSPackaging is a JavaScript framework that provides simple directives for defining, loading & importing uniquely named packages of JavaScript modules. UPDATE: This project now exists as Ajile at http://sf.net/projects/ajile & http://ajile.iskitz.com.1 weekly downloads
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Advanced learning tool to assist students in improving there Java programming skills. Includes real-time checking of user answers to exercises set through the web-based UI, which is powered by PHP, Java and mySQL.
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LAPdog - The Linux Acceptance Project
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Friday, July 29, 2016FOH Engineer Tomas Wolfe Chooses Roland M-5000 Console For The NeighbourhoodPSW Staff 07/29/16 02:08 PM,Front of house engineer Tomas Wolfe utilized a Roland M-5000 OHRCA digital console on both the U.S. and European legs of the recent tour by The Neighbourhood. Wolfe, a native of Los Angeles, worked his way up from the bottom in live audio, beginning with stints at Sunset Boulevard music venues such as the Viper Room and the Roxy Theatre. Two years on the L.A. club circuit led to touring with a series of artists, including BØRNS, Belly, Everlast, the… View this post
Filed in: AV • Live Sound • News • AV • Concert • Consoles • Digital • Engineer • Mixer • Sound Reinforcement
The Right StuffJonah Altrove 07/29/16 10:41 AM,A few years ago while working as an A1 for a local production company, I took a call from a young married couple who held an annual jam band festival on their property. In previous years the rig had been a potluck of cobbled-together "prosumer" gear, but they were seeking to bring in professional production to deal with bigger acts and a larger draw. For an expected attendance of 6,000, they requested side-by-side dual stages (plus the accompanying dual roof… View this post
Filed in: AV • Live Sound • Feature • Blog • Study Hall • AV • Business • Education • Engineer • Sound Reinforcement • Technician
Hal Leonard Publishes The Music Producer's Handbook, Second EditionPSW Staff 07/29/16 10:05 AM,The Music Producer's Handbook, Second Edition has just been published by Hal Leonard, revealing secrets to becoming a music producer and producing projects in any genre of music. Among the topics covered are the producer's multiple responsibilities and all the elements involved in a typical production, including budgeting, contracts, selecting the studio and engineer, hiring session musicians, and even getting paid. The Music Producer's Handbook, Second Edition also covers the true mechanics of production, from analyzing, troubleshooting, and fixing a… View this post
Filed in: Recording • News • Training • Business • Digital Audio Workstations • Engineer • Processor • Software • Studio • Technician
NEXO STM For Audi Open-Air Classical Festival In GermanyPSW Staff 07/29/16 09:36 AM,NEXO STM Series modular line arrays were deployed to serve a classical music audience at the Audi Art Experience in Germany. Rental company SHOWEM Veranstaltungstechnik GmbH presented a unique design of the modular system, including the largest deployment of STM M28 compact main modules to date, 48 in total. More than 20,000 people gathered together in the Klenzepark, the heart of the city of Ingolstadt for this year's classic concert. The main event featured the KlangVerwaltung Orchestra, conducted by Kent… View this post
Filed in: AV • Live Sound • News • AV • Concert • Line Array • Loudspeaker • Sound Reinforcement • Subwoofer
AES Sound Field Control Conference 2016 Breaks New GroundPSW Staff 07/29/16 09:16 AM,More than 90 delegates gathered in mid-June at the University of Surrey, Guildford, UK for the second international conference on Sound Field Control presented by the Audio Engineering Society (AES), where new ground was broken in the understanding of the active management of audio delivered within an acoustical environment. Over 30 papers were presented on topics ranging from sound zones through higher order ambisonics, mode matching and psychoacoustics, to sound field control theories, microphone arrays and array transducers. At the… View this post
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Production Sound Mixer Rashad Omar Utilizing DPA Microphones For Dialogue On High-Profile FilmsPSW Staff 07/29/16 08:56 AM,Production sound mixer and recordist Rashad Omar has been utilizing DPA d:screet minature microphones and d:dicate recording mics to capture dialogue for a number of high-profile films, including American Honey, which recently won the 2016 Jury prize at the Cannes Film Festival. Directed by Andrea Arnold and starring Sasha Lane, Shia LaBeouf and Riley Keough, the film is set in the American Midwest and tells the story of a teenage girl who joins a traveling magazine sales crew. A great… View this post
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Adamson E-Series Kicks Off Li Ronghao Tour In ShanghaiPSW Staff 07/29/16 08:26 AM,Popular Chinese pop singer, music producer and professional guitarist and Li Ronghao recently kicked off his latest tour with a concert at the Mercedes Benz Arena in Shanghai, reinforced by an Adamson Systems E-Series system provided by Beijing Global Visual Culture Development. "We're using a combination of 36 E15, 42 E12, 16 E218 and 18 E219 on the tour," explains Beijing Global Owner Li Ran. "This package provides the flexibility to cover virtually any venue we are dealing with." On… View this post
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64 Audio Launches apex And tia In-Ear Monitor Technologies (Includes Video)PSW Staff 07/29/16 08:00 AM,64 Audio has introduced apex in-ear monitoring technology, as well as tia, a tubeless in-ear system. apex stands for "air pressure exchange," incorporating a pneumatically interactive vent designed to relieve pneumatic pressure (air pressure) caused by miniature loudspeakers placed in a sealed ear canal. By reducing the pressure, sound quality is enhanced and a wider stereo image is created. In addition, instrument separation is more distinguishable. The technology is available in the apex-m20 module, specifically designed to fit the proprietary… View this post
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Thursday, July 28, 2016The Live Sound Loudspeaker Demo Is Coming To Louisville This SeptemberPSW Staff 07/28/16 09:22 PM,The Live Sound International (LSI) Loudspeaker Demo — a.k.a., "The Ultimate Audio Experience" — is coming to Louisville this September, presenting the unique opportunity to listen to, evaluate, and compare at least 24 top professional loudspeaker systems in a controlled listening environment. The event, which has attracted more than 3,000 attendees over the past several years, will be held in an exhibit hall in conjunction with this year's WFX (Worship Facilities Expo) at the Kentucky Exposition Center on September 21-22.… View this post
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MUSIC Appoints Goldsmith Sales & Marketing As Rep For Northwest U.S. TerritoryPSW Staff 07/28/16 02:53 PM,MUSIC has announced the appointment of Goldsmith Sales & Marketing as its manufacturer representative firm for the Northwest U.S. territory states of Washington, Oregon, Idaho and Alaska. Specifically, the firm will be promoting MUSIC's Enterprise Division brands, including MIDAS, KLARK TEKNIK, LAB GRUPPEN, LAKE, TANNOY and TURBOSOUND. The announcement was made by Chris Wildfoerster, senior director of Touchpoint Americas for the Enterprise Division. "Goldsmith Sales & Marketing has earned great respect from channel partners and consultants," states Wildfoerster. "MUSIC looks… View this post
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(HoS) Atmosphere3 - Creep
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pretty dark
awww no ones showing you love man, lol, well they do me the same way man, i'll show you love, it's pretty dark, it kinda reminds me of some silent hill type music but it's not enuff crazy shit going on to quilfie.... it's 3:46 in the am right now as i'm typeing this and this is very relaxing - think i'll go to sleep right now, it's a nice submission doh, nothing really bad to say about it, it does quite well :-) - good job homie...
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Yay! Best news all day
You probably already know this, but you can stream any TAL ever made, for free from their website.
And it's PRI, not NPR. Sorry, I worked briefly in public radio...
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text:
Truck flips, blocking I-26
Published: Saturday, March 20, 2010 at 4:30 a.m.
Last Modified: Friday, March 19, 2010 at 11:46 p.m.
SALUDA -- A tractor-trailer driver lost control on I-26 East, careened into a guard rail and smashed into an embankment, causing both the cab and trailer to flip onto its side Friday.
The accident happened near the 61 mile marker on the Saluda Grade.
The driver, James Pender of Lincolnton, suffered head trauma and was flown to Spartanburg Regional Hospital, according to Bob Johnson, manager of marketing and logistics at U.S. Tire Recycling. Pender was driving for the company, hauling hundreds of tires.
"Police, emergency people, state employees (NCDOT) and the wrecker service did a very professional job," Johnson said.
The one-vehicle accident occurred at 10:30 a.m. and closed both lanes of traffic. Shortly after crews arrived to the scene, one lane of traffic was open. However, both lanes were once again shut down momentarily as a front-loader tractor moved the mangled cab onto a flatbed truck, backing traffic up for 3 miles around 3:45 p.m.
Stuck in the traffic were several high school buses headed to the Wolverine Invitational track meet at Polk County High School.
The meet, which was supposed to start around 4:30 p.m., was delayed an hour-and-a-half due to at least four team buses being late because of the traffic backup.
Hazmat crews cleaned fuel and other fluids that poured into the culvert along the interstate as workers collected the tires and loaded them into another truck from the Concord-based recycling company.
Pender's condition was not released.
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text:
Gotabhaya Rajapaksa
|Gotabaya Rajapaksa
ගෝඨාභය රාජපක්ෂ
|Permanent Secretary, Ministry of Defence
|In office
November 2005 – January 2015
|Mahinda Rajapaksa
|Prime Minister
|Ratnasiri Wickremanayake
D. M. Jayaratne
|Preceded by
|Major General
Asoka Jayawardena
|Succeeded by
|B.M.U.D. Basnayake
|Personal details
|Nandasena Gotabaya Rajapaksa
June 20, 1949
Matara, Sri Lanka
|Sri Lanka
United State of America
|Sri Lankan
|Anoma Rajapaksa
|Mahinda Rajapaksa (brother)
Basil Rajapaksa (brother)
Chamal Rajapaksa (brother)
|Manoj Rajapaksa
|Alma mater
|Ananda College
|Military service
|Sri Lanka
|Sri Lankan Army
|Years of service
|Lieutenant Colonel
|Gajaba Regiment
|1st Gajaba Regiment
General Sir John Kotelawala Defence University
|Eelam War I
Eelam War II
Eelam War IV
Rana Sura Padakkama
Doctor of Letters (Honoris Causa), UoC
Lieutenant Colonel Nandasena Gotabaya Rajapaksa, RWP, RSP, psc, GR (Sinhalese: ගෝඨාභය රාජපක්ෂ) (born 20 June 1949) is a retired officer of the Sri Lanka Army, a former Secretary to Ministry of Defence and Urban Development of Sri Lanka. After serving through the early parts of the country's civil war with Tamil Tiger rebels, he retired from the Army in 1992. With the election of his brother Mahinda Rajapaksa as President of Sri Lanka, Gotabaya Rajapaksa was appointed Defence Secretary in November 2005.
As Defence Secretary, Rajapaksa played a key role towards the successes achieved by the Sri Lankan Military in defeating the Tamil Tigers and ending Sri Lanka's 26-year-long civil war in May 2009 with the political leadership of President Mahinda Rajapaksa and the military strategy of General Sarath Fonseka.[1][2][3][4][5][6][7] He was also one of the top targets of the Tamil Tigers, and survived an assassination attempt in December 2006 by a Tamil Tiger suicide bomber.[8]
- 1 Early life
- 2 Military career
- 3 Secretary to the Ministry of Defence
- 4 See also
- 5 References
- 6 Further reading
- 7 External links
Early life[edit]
Gotabaya Rajapaksa was born in Palatuwa in the Matara District,[citation needed] as the 5th of nine siblings and brought up in Weerakatiya in the southern rural district of Hambantota.[citation needed] According to a writer called Narada Karunthilaka, the name Gotabaya means "Abhaya, the Giant" or "Chinna Kotta" in the Jat language.[9] He hails from a well-known political family in Sri Lanka. His father, D. A. Rajapaksa, was a prominent politician, independence agitator, Member of Parliament, Deputy Speaker and Cabinet Minister of Agriculture and Land in Wijeyananda Dahanayake's government. His elder brother, Mahinda Rajapaksa was first elected to parliament as a member of the Sri Lanka Freedom Party at the age of 24 in 1970, who gradually rose through the party ranks becoming the Leader of the Opposition in 2001, Prime Minister in 2004 and the President of Sri Lanka in 2005. Two of his other elder brothers, Chamal Rajapaksa and Basil Rajapaksa, are also in the politics being current Members of Parliament.
He obtained his primary and secondary education at Ananda College, Colombo.[10] He has also obtained a Postgraduate degree in Information Technology from the University of Colombo in 1992.[11]
Military career[edit]
Rajapaksa joined the Sri Lanka Army as a Cadet Officer on April 26, 1971, when Sri Lanka was still a dominion of the British Commonwealth. He was commissioned as a Second Lieutenant on May 25, 1972 and given his first command as an officer in the Ceylon Signals Corps after his basic training at the Army Training Centre, Diyatalawa. Thereafter he served with the Sri Lanka Sinha Regiment and the Rajarata Rifles before being transferred to the Gajaba Regiment upon its formation in 1983 with the amalgamation of the Rajarata Rifles and Vijayabahu Infantry Regiment.[12]
During his 20 years of military service, Rajapaksa has received awards for gallantry from three Presidents of Sri Lanka, J.R. Jayewardene, Ranasinghe Premadasa and D.B. Wijetunga.[13] He had followed signal young officers course in school of signals at Rawalpindi; Infantry company commanders course in Queta; Jungle warfare and counter insurgency course in Assam; Command and Staff course at Defence Services Staff College in Welington; and Advanced Infantry Officers course at Fort Benning.
He rose up the ranks in the military, serving as the second in command of the 1st Gajaba Regiment and latter commanding the 1GR from 1983 to 1990. He served in the battlefronts of Jaffna, participating in Operation Liberation, the offensive mounted to liberate Vadamarachi from LTTE in 1987. He also commanded the same battalion in Operation "Strike Hard" and Operation "Thrivida Balaya" in 1990. He was promoted to Lieutenant Colonel, and served as the Commandant of the Kotelawala Defence Academy at the time of his retirement in 1992. He subsequently migrated to the United States. Also, he worked in Loyola Law School[14] in Los Angeles, U.S. as a Systems Integrator and Unix Solaris Administrator.[15]
Secretary to the Ministry of Defence[edit]
In order to assist his brother's Presidential election campaign, Rajapaksa returned to Sri Lanka from the United States in 2005 .
Gotabaya Rajapaksa was appointed to the post of Permanent Secretary of the Ministry of Defence in November 2005 by newly elected President Mahinda Rajapaksa. In this capacity he oversaw the military operation which eventually defeated the LTTE in May 2009.
At the public acclaim as a "war hero," he was conferred Doctor of Letters by the University of Colombo on September 6, 2009.[16][17] He is also the secretary to the Urban Development, with the help of the army he undertakes huge projects.
Attempted assassination[edit]
On December 1, 2006, at approximately 10:35 an assassin attempted to drive an explosive laden auto-rikshaw into Rajapaksa's motorcade as it traveled through Kollupitiya, Colombo. The Sri Lanka Army Commandos guarding him obstructed the vehicle carrying the explosives before it reached Rajapakse's vehicle and two commandos were instantly killed. Rajapaksa escaped unhurt.[8] The LTTE were blamed for the attack.[8]
Criticism of United Nations and western countries[edit]
In June 2007, Rajapaksa was severely critical of the United Nations (UN) and of western governments. He accused the UN of having been infiltrated by terrorists "for 30 years or so", and as a result the UN was fed incorrect information. He also alleged that Britain and the EU were bullying Sri Lanka, and concluded that Sri Lanka "does not need them", and that they don't provide any significant amount of aid to the country.[18]
War Crimes[edit]
On 3 February 2009, Defense Secretary Gotabhaya Rajapaksa stated to the international media (in reference to the only hospital in the war front being shelled), that everything is a legitimate target if it's not within the safe zone the government has created and that all persons subject to attack by the armed forces were legitimate LTTE targets as there are no independent observers, only LTTE sympathizers, radio announcements were made and movement of civilians started a month and a half ago.[19][20]
As per wikileaks, General Sarath Fonseka who lead the war against LTTE had accused Defense Secretary Gotabhaya Rajapaksa of ordering at the end of the war the shooting of any LTTE leaders who might try to surrender under flags of truce. But Mr. Gotabhaya alleged to have threaten to execute Mr. Fonseka if he had spilled any war secrets.[21] [22] [23]
In an interview on the Sri Lanka TV channel adaderana on March 16, 2015, Mr Rajapaksa stated that he is a citizen of the United States but cannot travel to the United States because of alleged war crimes charges against him.[24]
Karuna link[edit]
Gotabhaya is credited with using the Karuna faction effectively during the war to defeat the LTTE. The former LTTE commander Vinayagamoorthi Muralitharan, better known as Colonel Karuna, told British authorities that Rajapaksa was instrumental in arranging for him to be issued with a false diplomatic passport so that he could flee to Britain in September 2007. These allegations were denied by the Sri Lankan Foreign Minister Rohitha Bogollagama at the time,[25] and later by Rajapaksa.
Assassination of Nadarajah Raviraj[edit]
Nadarajah Raviraj, a well-known human-rights lawyer and a parliamentarian, was shot and killed in Colombo on 10 November 2006. At a magisterial hearing in Colombo on 26 February 2016, Liyanarachchi Abeyrathna, a former policer officer attached to the State Intelligence Agency, stated that Gotabhaya Rajapaksa paid Rs. 50 million to a terror organization led by Colonel Karuna to murder Mr. Raviraj.[26][27]
Relationship with the media[edit]
Rajapaksa has been accused of threatening journalists on several occasions, including telling two journalists attached to the state-owned Lake House Publications that unless they stop criticising the armed forces "what will happen to you is beyond my control". When asked by the two journalists if he was threatening them, he replied "I am definitely not threatening your lives. Our services are appreciated by 99 per cent of the people. They love the Army Commander (General Sarath Fonseka) and the Army. There are Sri Lankan patriots who love us do and will do what is required if necessary."[28] In April 2007 he was accused of allegedly calling the Editor of the Daily Mirror Champika Liyanaarachchi and threatening her, saying that she would escape reprisals only if she resigned.[29] He was also accused for threatening to "exterminate" the Daily Mirror journalist Uditha Jayasinghe for writing articles about the plight of civilian war casualties.[30][31][32]
A December 5, 2008 story from The New York Times quoted his news reporting position as "he insists that journalists should not be allowed to report anything that demoralizes the war effort".[33]
In the editorial titled A brother out of control (August 16, 2011), The Hindu raised the observation, "President Rajapaksa would be well advised to distance himself swiftly from his brother's stream-of-consciousness on sensitive issues that are not his business. This includes an outrageous comment that because a Tamil woman, an "LTTE cadre" who was a British national, interviewed in the Channel 4 documentary was "so attractive" but had been neither raped nor killed by Sri Lankan soldiers, the allegation of sexual assault by soldiers could not be true. For this statement alone, Mr. Gotabaya Rajapaksa must be taken to task."[34]
Political Abductions & White Van Incidents[edit]
As reported by The Sunday Leader, Major General Prasad Samarasinghe the former Military Spokesman and Director, Directorate of Media in the Army, has been passing highly sensitive information to the US Embassy in Colombo on abductions. Many of those abducted were believed to have been individuals who had fallen foul of the Rajapaksa trio, Mahinda, Basil and Gotabhaya. During her visit to the country, United Nations Human Rights Commissioner Navanetham Pillay expressed her disappointment over "white van" related disappearances reported in Colombo, and other parts of the country, which were not covered by the Commission of Inquiry on Disappearances set up by the government.[35][36][37]
Deceased MP Sripathi Sooriyarachchi, assassinated journalist Lasantha Wickrematunge and others had accused Rajapaksa of corruption since 2006.[38][39] In 2015 Interpol provided further evidence to the Sri Lankan government on corrupt military procurements.[40][41][42][43] In March 2015 a Sri Lankan court imposed a travel ban on Rajapaksa over allegations he used a commercial floating armory as a private arsenal.[44][45]
From 2015[edit]
After the defeat of Mahinda in the 2015 Presidential elections, he was replaced as Secretary of Defence by B.M.U.D. Basnayake a day after the new President was sworn in.[46]
See also[edit]
- Eelam War IV
- List of attacks attributed to the LTTE
- List of Sri Lankan non-career Permanent Secretaries
- ^ President Maithripala Sirisena's Address to the Nation at the 67th Independence Day celebrations, NEWS.LK, Retrieved 29-07-15
- ^ Mehta, Maj. Gen. (Retd) Raj K. Lost Victory: The Rise & Fall of LTTE Supremo, V. Prabhakaran. Pentagon Press. pp. 13, 118, 330–338, 360–8, 377, 380, 401–11. ISBN 978-8182744431.
- ^ Review - Gota's War -Final- revised By Major General (Retired) Lalin Fernando, Asian Tribune, Retrieved 18 May 2015
- ^ Mi 24s role in eelam war IV emphasised by Shamindra Ferdinando, The Island, Retrieved 17 May 2015
- ^ Refurbishing the Armed Forces By Tissa Ravindra Perera, The Nation, Retrieved 18 May 2015
- ^ SF denies claim of 40,000 civilian deaths in Vanni, says his men paid heavy price by not using heavy weapons By Shamindra Ferdinando, The Island, Retrieved 21 May 2015
- ^ Fonseka to become first Sri Lankan Field Marshal, Khaleej Times (UAE), Retrieved 29 March 2015
- ^ a b c "CHRONOLOGY-Attacks blamed on Sri Lanka's Tamil Tigers". Reuters. 2008-01-08.
- ^ www.toplankanews.com[dead link]
- ^ Meteoric rise of Ananda's patriotic sons, Rasika SOMARATNA, Daily News, Retrieved 4 May 2015
- ^ A legend of our times – Opinion. Defence.lk. Retrieved on 2012-07-09.
- ^ Who are the Real Traitors ?. www.defence.lk (2008-12-30). Retrieved on 2012-07-09.
- ^ T.B. Singalaxana (2007-07-19). Politics, Thoppigala and rhetoric at the Wayback Machine (archived September 30, 2007). Daily Mirror
- ^ The Honourable Gotabaya Rajapaksa
- ^ 1 Gotabaya Rajapaksha-Talk at "Tech Colloquium" organized by Microsoft on YouTube. Retrieved on 2012-07-09.
- ^ Talk at the Cafe Spectator, The Sunday Times, Retrieved 29-07-2015
- ^ "Honorary degrees to the President and Gota". Daily Mirror. 2009-09-05. Retrieved 2009-11-03.
- ^ Buerk, Roland. (2007-06-12) South Asia | Sri Lanka accuses 'bullying' West. BBC News. Retrieved on 2012-07-09.
- ^ CNN-IBN (2009-02-03). "'Can't ensure civilians safety in LTTE area'". CNN IBN live. Retrieved 2015-04-05.
- ^ Ryan Goodman (2014-05-19). "Sri Lanka's Greatest War Criminal (Gotabaya) is a US Citizen: It's Time to Hold Him Accountable". Just Security. Retrieved 2015-04-05.
- ^ "Gotabaya Rajapaksa should be tried for war crimes". Frontierindia.net. 2013-04-14. Retrieved 2013-11-01.
- ^ "Fonseka will be hanged if he spills war secrets: Lanka". Times Of India. 2010-06-07. Retrieved 2013-11-01.
- ^ Jason Burke, South Asia correspondent (2011-11-18). "Former Sri Lankan army chief convicted for war crimes claim | World news". The Guardian. Retrieved 2013-11-01.
- ^ "Gotabaya reveals why he can't go back to the US.". www.youtube.com. 2015-03-16.
- ^ "Gotabaya 'gave me passport'". BBC News. 2008-01-25. Retrieved 2009-11-03.
- ^ "Gota Paid Karuna Faction 50 Million To Kill Raviraj". Colombo Telegraph. 2016-02-27. Retrieved 2016-02-27.
- ^ "Gotabaya has given 50 million for the Raviraj killers". Lanka News Web. 2016-02-27. Retrieved 2016-02-27.
- ^ Sunday Times – Death bells toll for the free media. Sundaytimes.lk (2008-06-01). Retrieved on 2012-07-09.
- ^ :: Daily Mirror on the Web – Front Page ::[dead link]
- ^ "FMM protests Gotabhaya's 'threats'". BBC Sinhala. 5 May 2008.
- ^ "Sri Lanka - Annual report 2008". Reporters Without Borders.
- ^ "Britain exposes damn lie by the state media". The Daily Mirror (Sri Lanka). 21 April 2007.
- ^ Somini Sengupta (2008-12-05). Sri Lankan Army Is Pushing for End to 25-Year War Against the Tamil Rebels, The New York Times.
- ^ "A brother out of control". The Hindu. 2011-08-16. Retrieved 20 August 2011.
- ^ "A Top Sri Lankan Major General, The US And Gotabhaya Rajapaksa". The Sunday Leader. Retrieved 2013-11-01.
- ^ "Video: Signs of Sri Lanka moving towards authoritarianism-Pillay | Breaking News". Dailymirror.lk. 2013-08-31. Retrieved 2013-11-01.
- ^ "Opening remarks by UN High Commissioner for Human Rights Navi Pillay at a press conference during her mission to Sri Lanka Colombo". Office of the United Nations High Commissioner for Human Rights. 31 August 2013. Retrieved 6 September 2015.
- ^ Rajapaksa's brother probed over killings, aljazeera, Retrieved 2015-06-06
- ^ Academic Exposing Corrupt War Procurement Tender Process Threatened By Rajapaksa Government, Colombo Telegraph. Retrieved 2015-06-06
- ^ MiG Deal: Rajapaksas Paid US$ 10 Million To A Ghost Company: "No Company Called Bellimissa" – Interpol Confirmed, Colombo Telegraph. Retrieved 2015-05-11.
- ^ FCID questions Gotabaya, adaderana. Retrieved 2015-06-06
- ^ A Trembling Gotabaya Appears Before FCID, Colombo Telegraph. Retrieved 2015-06-06
- ^ Gota's MiG Deal: Investigation Finds Secret Bank Account In British Virgin Islands, Colombo Telegraph. Retrieved 2015-06-06
- ^ "Sri Lankan court imposes travel ban on former defence secretary Rajapaksa". The Guardian. Retrieved 22 July 2015.
- ^ "Sri Lanka travel ban for Gotabhaya Rajapaksa over arms ship". BBC. Retrieved 22 July 2015.
- ^ "B.M.U.D. Basnayake appointed new Defence Secretary". Ada Derana. 10 January 2015. Retrieved 31 March 2016.
Further reading[edit]
- Army, Sri Lanka. (1st Edition – October 1999). "50 YEARS ON" – 1949–1999, Sri Lanka Army. ISBN 955-8089-02-8
- The Broken Palmyra – The Tamil Crisis in Sri Lanka: An Inside Account. The Sri Lanka Studies Institute, Claremont 1990. By Hoole, R., Somasundaram, D., Sritharan K., and Rajini Thiranagama. (Also available online.)
- War and Peace in Sri Lanka: With a Post-Accord Report From Jaffna. ISBN 955-26-0001-4/ISBN 978-955-26-0001-2, Institute of Fundamental Studies, Sri Lanka; 1 edition (October 1, 1987), By Rohan Gunaratna.


text:
Monday, 05 Mar 2012 11:36 AM
CUPERTINO, Calif. — More than 25 billion apps have been downloaded from Apple's app store.
The downloads came from more than 315 million iPhones, iPads and iPod touches, according to a news released from the Cupertino-based Apple on Monday. The app store has more than 550,000 apps, some of which are free.
Chunli Fu of Qingdao, China, downloaded the 25 billionth app and won a $10,000 iTunes gift card.
The app store launched four years ago. Apple says the store has paid out more than $4 billion to developers.
© Copyright 2016 The Associated Press. All rights reserved. This material may not be published, broadcast, rewritten or redistributed.
Newsmax, Moneynews, Newsmax Health, and Independent. American. are registered trademarks of Newsmax Media, Inc. Newsmax TV, and Newsmax World are trademarks of Newsmax Media, Inc.


text:
In its bid to bolster the biotechnology sector
and improve the quality of European food, the Commission has
launched three new agrofood research projects and a training
programme to help scientists gain some entrepreneurial know-how.
The European Commission has pumped €43
million into three new agrofood research projects. "Meeting
consumers' expectations for good and safe food, while at the
same time ensuring sustainable agriculture and fostering a
competitive industry, requires ambitious R&D efforts and
strong co-operation at European level," Research Commissioner
Philippe Busquin said during the launch of the new projects
earlier this month.
He added that these projects are the first under the 15-month-old
Sixth Framework Programme (FP6) to take the EU's comprehensive
'farm-to-fork' approach which "opens new
perspectives for producing food that respond to environmental,
health and competitiveness objectives".
The Safe Foods project aims to improve consumer protection
by formulating an integrated food safety risk analysis approach.
The NuGO network seeks to integrate and develop nutritional
genomics in Europe. Welfare Quality strives to integrate animal
welfare into the food production chain .
Learning the trade
In addition to providing its citizens with the tastiest and
safest food possible, Europe aims to become the world's
most competitive knowledge-based economy by 2010. "Growth
and job creation in key sectors such as life sciences and
biotechnology are mainly driven by new companies," Mr
Busquin has observed.
This involves promoting entrepreneurship among European scientists
and researchers. Towards that end, the Commission has just
launched a fourth series of its successful Biobiz workshops.
"One of the barriers for researchers who would like
to create their own company is the lack of knowledge and training
in management and finance… Biobiz helps fill this gap
and meet the entrepreneurial challenge in the biotechnology
sector," the Research Commissioner explained.
Biobiz will hold some 30 workshops in Europe and the USA over
the next three years. Since its inception in 1998, around
350 prospective entrepreneurs have already attended such events.
The initiative has helped to create 40 new biotech companies
Source: EU sources
More Information:
Commission press release
Biobiz workshops
Information on life sciences, genomics and biotechnology for health under FP6


text:
Reload the page and try again.
Free printing is wonderful and there is availability.
I didn't need a laptop until my second year when the workload was getting heavier. There are plenty of computer facilities.
We don't have as much in terms of computer labs. Around semester's end when everyone has projects and papers due, the labs are packed. I would definitely recommend bringing a computer. I have friends that do not have laptops, and regret it. Printing is completely FREE of charge, but copies are not. The network reliability/speed is okay.. However, wi-fi is not available in the dorms. You need to bring your own router or ethernet cable. There are printers to use on campus but beware, the ones in the library can only print documents etc from Microsoft Word programs. During crunch times, such as right before classes, lines for printers can be a little long but not too bad. I personally bought my own printer to make my life easier.
It's a great college, it's just a very small town.
The computer labs are usually pretty empty and if they are full, ask a librarian. There are computers seemingly everywhere around the library and under the commons. Everyone is pretty open about using computers and very rarely do you find someone messing around on them. Also, having your own laptop it great. The college has wireless that can be tapped into everywhere.
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Ripon is quite good about providing for its students when it comes to computers or Internet access. There are several labs around campus: Kemper, the Collaborative Learning Center (CLC), the library, as well as labs in academic departments. Wherever there is a printer on campus, printing is free! There are printers in all of the labs, as well. The only color printer on campus is in the library, and in order to print there, you must inform the person working at the desk that you've printed something and then he or she grabs it for you.
There is Wi-Fi in most common places on campus such as the Commons, the Union/Pub, the library, and most academic buildings. There are also lounges in each residence hall that have Wi-Fi. So while you might not be able to access Wi-Fi from your room, there is a lounge in each res hall that has it. There are also Ethernet jacks in each room for each student. Students can purchase an Ethernet cable before coming to campus or on campus in the college bookstore, and then plug their laptop or computer into the cable that plugs in to the wall and receive Internet that way, as well. While a majority of students has laptops on campus, it is not a necessity. Some have desktops or nothing at all, and they just rely on computers around campus for any Internet or computer needs.
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