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Claims of extreme survival of DNA have emphasized the need for reliable models of DNA degradation through time. By analysing mitochondrial DNA (mtDNA) from 158 radiocarbon-dated bones of the extinct New Zealand moa, we confirm empirically a long-hypothesized exponential decay relationship. The average DNA half-life within this geographically constrained fossil assemblage was estimated to be 521 years for a 242 bp mtDNA sequence, corresponding to a per nucleotide fragmentation rate (k) of 5.50 × 10–6 per year. With an effective burial temperature of 13.1°C, the rate is almost 400 times slower than predicted from published kinetic data of in vitro DNA depurination at pH 5. Although best described by an exponential model (R2 = 0.39), considerable sample-to-sample variance in DNA preservation could not be accounted for by geologic age. This variation likely derives from differences in taphonomy and bone diagenesis, which have confounded previous, less spatially constrained attempts to study DNA decay kinetics. Lastly, by calculating DNA fragmentation rates on Illumina HiSeq data, we show that nuclear DNA has degraded at least twice as fast as mtDNA. These results provide a baseline for predicting long-term DNA survival in bone.
1. Introduction
Although early-1990s claims of DNA recovered from million year-old fossils [1–4] are now widely regarded as modern contaminants [5–13], the kinetics of long-term post-mortem DNA decay is still poorly understood. There is a lack of empirical data on which to estimate the rate of in situ DNA fragmentation. Because the field of ancient DNA (aDNA) has recently entered the era of whole-genome profiling [14–17], which is dependent on samples of exceptional preservation, understanding the nature and rate of DNA decay is as pertinent as ever, for both predictive and authentication purposes.
DNA has limited chemical stability and decays without the enzymatic repair mechanisms of living cells [18]. Following cell death, nucleases start to cleave the DNA into fragments [19] and during decomposition, the DNA is digested by micro-organisms [18,20]. In determining long-term DNA decay, it is believed that hydrolysis of amino groups accelerates the loss of purine residues (depurination), resulting in strand cleavage [21,22]. This random DNA fragmentation generates a characteristic negative exponential correlation between DNA fragment length and number of molecules (figure 1a) [23,24,26,27].
Although other types of DNA damage occur, such as cross-links [28,29], the role of depurination in DNA fragmentation is well documented, and damage patterns from high-throughput sequencing data show that purines are overrepresented adjacent to strand breaks in ancient samples [30–33]. However, it is not known whether the rate of fragmentation can be regarded as constant through time, nor to what extent it varies between specimens from similar depositional environments. Indeed, a constant rate would imply that DNA decay follows first-order kinetics (i.e. DNA has a half-life; figure 1b), and hence that the age of a specimen can potentially work as a proxy for its DNA preservation and vice versa.
Despite many attempts [24,31,34–41], it has proved extremely difficult to demonstrate a general association between age and DNA preservation, probably because of variation in physical, chemical and biological factors such as taphonomy, fossil storage, oxygenation, microbial diagenesis, pH and ionic strength, and the presence of cations, humics and humates. Moreover, most studies are limited by small sample sizes and lack of individually dated specimens, leading to potential problems of mixed taphonomies. The absence of clear temporal trends in previous work suggests that the rate of DNA decay either fluctuates widely (hence not a rate per se), or that it takes a large, homogenous, and accurately dated sample, to overcome the 'noise' introduced by the aforementioned factors. In the light of many inconclusive results, a common refrain is that DNA decay rates cannot be predicted [42–44].
A few studies, however, have indicated a relationship between DNA preservation and post-mortem age. Bar et al. [45] showed that the length of amplifiable DNA fragments from human tissues (blood, muscle and spleen) decreased exponentially during the 20 days after a person's death, and Campos et al. [46] showed a rapid decrease in DNA content in cow bone after burial. Moreover, Adler et al. [26] showed a weak exponential correlation between DNA preservation and age, on the basis of 37 ancient human samples (mainly teeth). These samples had not all been individually dated, but the result is encouraging and provides impetus for more detailed investigations of long-term DNA fragmentation.
Establishing an association between age and preservation is not the only challenge. The rate of depurination is influenced by temperature, among other factors [22], which explains why the most extreme survival of DNA was documented in approximately 450–800 kyr ice cores [47]. Smith et al. [11,12] argued for the possibility of using the temperature-dependence of DNA fragmentation to normalize the samples and predict DNA survival. Such a relationship has been demonstrated for collagen, the most abundant protein in bone [48]. However, to establish a thermal age model for DNA, the first step is to confirm that long-term in situ DNA degradation can be described by a rate kinetic.
In an attempt to document a correlation between sample age and DNA preservation, we use a quantitative real-time PCR (qPCR) design to measure relative copy numbers of mitochondrial DNA (mtDNA) fragments from bones of the extinct New Zealand moa (Aves: Dinornithiformes). Aside from a large dataset (qPCR data from 158 radiocarbon-dated bones), our study differs from previous attempts by being highly localized. Using Holocene fossil tibiotarsi recovered from anoxic, limestone-buffered sediments at three adjacent (less than 5 km apart) fossil deposits (figure 2), we endeavour to provide a high level of sample homogeneity and minimize variables of climate and taphonomy. Also, by sampling only natural bone accumulations, we expect to minimize other complicating factors (e.g. butchering and cooking) that could impact on taphonomic processes [27,49].
Because the three fossil sites were excavated at different times (figure 2), the influence of storage times on DNA recovery can also be assessed. Pruvost et al. [50] argued that DNA degradation intensifies when a bone is removed from its deposition environment. The moa fossils analysed here were recovered at well-documented dates over an approximately 70 year period, allowing us to test the importance of post-excavation storage time.
The three main objectives of this study were: (i) to test whether long-term DNA decay follows first-order kinetics, thereby confirming the foundation for a predictive model; (ii) to estimate the long-term decay rate in bone at a given burial temperature and compare this rate with the predicted depurination rate from DNA in solution [21,22]; and (iii) to estimate the relative importance of storage time on DNA preservation in bone.
Lastly, following Deagle et al.'s [23] demonstration that frequency distributions of DNA fragment lengths from a sample of known age can be used to estimate the DNA decay rate, we argue that histograms based on the thousands-to-millions of reads generated from high-throughput sequencing platforms could represent a 'by-product' to investigate DNA fragmentation. We therefore apply the method to Illumina HiSeq 2000 data generated from two moa specimens, in order to achieve independent DNA decay-rate estimates, comparable with the qPCR results.
2. Material and methods
(a) Sampling, dating and storage
Sampling of 158 moa left tibiotarsi was conducted as described previously [51]. The material covered three adjacent sites, located within a 5 km radius in North Canterbury, South Island, New Zealand; Pyramid Valley (PV; n = 103), Bell Hill Vineyard (BHV; n = 47) and Rosslea (n = 8) (figure 2). Further information on samples and sites has been given previously [52,53] and is also listed in electronic supplementary material, table S1. The PV and BHV specimens were sampled from museum collections, whereas the Rosslea material was sampled less than 14 days after excavation in 2008 [53].
Bone gelatin was extracted by Isolytix Ltd, Dunedin, New Zealand, using standard protocols. Gelatin sample splits were sent for stable isotopic analysis (Environmental Isotopes Ltd, Sydney, Australia) and accelerator mass spectrometry (AMS) radiocarbon analysis. Radiocarbon ages were measured at the Rafter Radiocarbon Laboratory, GNS Science. Lower Hutt, New Zealand. Ages are reported as conventional radiocarbon ages, and as median calibrated ages and 95% probability distributions generated by the OxCal v. 4.1 (Oxford Radiocarbon Accelerator Unit) program. Sample quality was assessed using the C : N molar mass ratio, and stable isotopic ratios (δ15N, δ13C) in comparison with archive data, and was uniformly high (see the electronic supplementary material, table S1).
The moa bones used in this study had been excavated at different times (figure 2). The PV elements were recovered during excavations from 1939 to 1973 [54], BHV was excavated in 2001 [52] and Rosslea in 2008 [53]. To assess the potential DNA degradation effect of storage time, a multiple regression analysis (SPSS v. 19) was conducted between DNA preservation, calibrated 14C age and storage time. This analysis was conducted on the entire dataset (n = 158) and exclusively on 103 elements from PV that were excavated over a 35-year time frame, with excavation dates recorded for each bone.
(b) DNA extraction and qPCR
DNA was isolated from all samples at dedicated aDNA facilities (Murdoch University, Perth, Australia), using a standardized method described previously [52] and detailed in the electronic supplementary material, text S1. All DNA extractions were undertaken from 200 mg uniformly sampled and powdered bone. The DNA isolation protocol (molecular weight cut-off (MWCO) and Silica purification) was selected owing to its ability to generate DNA largely free of inhibitors.
DNA amplifications of the 262F/441R fragment (mtDNA control region, 242 bp including primers—see Bunce et al. [55]) were performed using SYBR-green detection chemistry on a BioRad My-IQ thermocycler. With a standardized level of detection, the recorded cycle threshold (CT) values represented the relative DNA preservation in each extract. To minimize effects of post-extraction storage time, qPCR was conducted immediately following DNA extraction. The quantitative response and general applicability of our qPCR set-up has been validated before based partly on the same extracts [51,56–58]. In addition, we conducted melt curve analyses for all PCR products, (see the electronic supplementary material, text S1 and figure S1), as well as 'spike-in' (see the electronic supplementary material, table S2), dilution- (see the electronic supplementary material, figure S3 and table S3) and PCR efficiency (see the electronic supplementary material, figure S3) experiments. This was done to ensure that amplification of the correct target was being monitored and to test for PCR inhibition (see the electronic supplementary material, text S1). In most instances (greater than 70%), the actual qPCR products were directly sequenced, confirming their species origin (data not shown) and specificity of the amplicons. Extracts displaying deviating melt curve diagnostics (owing to non-specific amplifications and/or dimer accumulation) were omitted from the analysis (ca 20 samples). We acknowledge that the relatively large (242 bp) amplicons, with primers that had to work on multiple species and across a large dynamic range of DNA yields, is not ideal in a qPCR set-up. All of our tests indicated, however, that the assay is sufficiently robust and reproducable for the intended purpose.
Stochastic variation between qPCR reactions is expected, in particular when amplifying low copy number DNA at CT values greater than 35. Results from 50 extracts in repeated qPCRs (same extract, but independent PCR mastermix and thermocycling run) were used to isolate this effect and estimate the variation introduced by stochastic qPCR effects. To test for consistency across DNA extracts, one moa bone was sampled and extracted three times, showing only minor deviations in CT (less than 1 cycle) between extracts at multiple levels of dilution (see the electronic supplementary material, table S4).
Preliminary data suggested that DNA from one of the four North Canterbury moa species (Pachyornis elephantopus) amplified less efficiently in the qPCR assay. We suspect that primer-binding-site mutations are the cause, and to prevent a systematic bias, data from this species were excluded.
(c) Estimating the rate of decay from qPCR data
The number of DNA copies in a PCR reaction follows an exponential increase per cycle. If one DNA extract produces a CT three cycles higher than another, then the former has eight times (23) less DNA (in an ideal situation with 100% amplification efficiency). To test for an association between DNA preservation and age, we calculated the relative number of template molecules in each sample according to the 'best' sample (Dinornis robustus, S39955, CT = 24.8), which was set to 100 per cent. Although some previous aDNA studies convert CT values into absolute starting copy numbers [24,59], others do not [46] as the accuracy and meaningfulness can sometimes be questioned [58]. For our purpose, the use of relative CT values was sufficient, as we were interested only in rates and not in absolute numbers.
Following Adler et al. [26], we tested (in SPSS v. 19) whether the observed DNA degradation through time would be better described by a linear, inverse, S-shaped or exponential decay model. An exponential decay relation could be estimated as: Nt = N0 × ek242t (Nt and N0 being the quantity at time t and 0, respectively, and k242 the decay constant for the entire 242 bp fragment). The average molecular half-life (t1/2 = (ln 2)/k242) could be calculated from the entire dataset and for each fossil site, respectively. A model of random fragmentation predicts that the relationship between decay rate and fragment length is a Poisson distribution (figure 1) [23]. The average per nucleotide rate of decay is therefore calculated by isolating k in: k242 = 1 – (e−k*242).
(d) Estimating the rate of decay from Illumina HiSeq data
DNA extracts of two South Island giant moa (D. robustus) fossils were selected for Illumina HiSeq 2000 sequencing (see the electronic supplementary material, text S1 for details). Only a fraction of 'shot-gun' sequencing data from an ancient bone is expected to represent DNA of the target species. Microbial DNA will constitute a proportion of the reads [60,61]. To retrieve sequences of endogenous moa DNA, only reads that mapped against an unpublished draft of the ostrich (Struthio camelus) genome (source tissue provided by San Diego Zoo and advance access to genomic data provided courtesy of Oliver Ryder, San Diego Zoo; Genome 10K; BGI, China) and the mtDNA genome of a D. robustus moa (GenBank: AY016013.1) [62] were used in the downstream analyses. The software BWA [63] was used in the mapping, using default settings. The resulting fragment length histograms of all mapped reads were used to assess DNA decay rates. According to a model of random fragmentation, the amount of amplifiable template should decline exponentially with increasing fragment size [23] (figure 1a). Log-transformed copy numbers therefore have a linear relationship with amplicon length, with the slope of the decline (λ) describing the probability of a bond in the DNA backbone being cleaved [23]. Lambda can be converted to a damage rate when sample age is known (figure 1b).
3. Results and discussion
(a) Estimating the half-life of mtDNA in bone
The calibrated radiocarbon ages of the 158 fossils documented a Mid- to Late Holocene accumulation, ranging from 602 to 7839 BP, calibrated calendar years before present ('present' defined as CE 1950; figure 2; electronic supplementary material, table S1). When correlating relative DNA copy number with age, a linear (R2 = 0.002, p = 0.604), inverse (R2 = 0.002, p = 0.598) and S-shaped (R2 = 0.229, p < 0.001) model proved inferior to an exponential relationship (R2 = 0.386, p < 0.001; figure 3). Hence, the qPCR results demonstrated that the number of template mtDNA molecules in the fossils was best described with first-order decay kinetics (see the electronic supplementary material, table S4 for individual CT values). The equation for the fitted regression was: 3.1with Nt being the proportion of molecules left at the time t (years BP), compared with the best sample. Solving the equation yielded a molecular half-life of 521 years for the targeted 242 bp mtDNA fragment, and assuming a Poisson distribution [23], the per site decay rate (k) was 5.50 × 10–6 per year.
As noted, several studies have attempted to investigate DNA degradation kinetics using samples of different ages. Here, we show that by using a controlled experimental set-up, with samples normalized for ambient temperature, an exponential relationship between sample age and DNA preservation can be demonstrated. This supports the view that random DNA fragmentation, likely caused by depurination, applies not only to short-term degradation of DNA in a solution [21,22] but also to ancient DNA in fossil bone.
(b) Comparison with Lindahl & Nyberg's [22] results
The rate constant (k) depends on absolute temperature (T) according to the relationship: 3.2where A is the pre-exponential factor, Ea is the activation energy and R the gas constant. On the basis of the Arrhenius plot in Lindahl & Nyberg [22], showing the temperature-dependency of depurination at pH 5, we estimated k to 2.11 × 10–3 per site per year (see the electronic supplementary material, text S2), which is 384 times faster than the rate estimated from the moa data (figure 4).
Assuming that long-term DNA fragmentation happens primarily as a result of depurination, this discrepancy is probably related to pH and access to free water. Even if the burial environment is slightly acidic, sacrificial dissolution of microcrystalline carbonated bioapatite will likely act as buffer within the bone. On the basis of the measurements in Berna et al. [64], we used pH 7.5 as an appropriate value for bone (see the electronic supplementary material, Text S2). From Lindahl & Nyberg [22], we find that k at a pH of 7.5 is ca 73 times slower than the same at pH 5 (see the electronic supplementary material, Text S2), but still ca five times faster than the rate observed for moa mtDNA (figure 4). Given that the same authors also reported a twofold rate decrease in the presence of apatite, as in bone (cited in [18]), the rate expected from Lindahl & Nyberg's [22] in vitro results is perhaps only roughly twice as fast, as the rate we observe for the moa qPCR estimate.
From equation (2.2), it is clear that both the pre-exponential factor (A) and the activation energy (Ea) affect the decay rate (k), but Lindahl & Nyberg [22] showed similar values of Ea at pH 5 and 6. We therefore assume that the estimated Ea of 127 kJ mol−1 also applies to pH 7.5. With this Ea value, and an observed average k for moa mtDNA of 5.5 × 10–6 per site per year, and an estimated fossil burial temperature T = 13.1°C or 286.25 K (see the electronic supplementary material, text S3), the pre-exponential factor A is determined to e41.2. The average decay rate (per site per year) for mtDNA in moa bone is therefore related to temperature as follows (figure 4): 3.3
This is arguably the best available approximation for the temperature-dependency of long-term decay of mtDNA in bone, and it should be used as a guide for future work on DNA decay kinetics. We note that Lindahl & Nyberg [22] tested only the temperature dependency of depurination between 45°C and 80°C (318.15–353.15 K), and it remains to be confirmed that the relationship in their Arrhenius plot can be projected to include more typical burial temperatures, as seen in table 1. Furthermore, we note that our model of mtDNA decay in bone does not account for a potential initial post-mortem phase of faster DNA decay, facilitated by nuclease activity rather than depurination [23,31,65]. With these caveats in place, table 1 shows our predicted half-lives, and average fragment lengths of mtDNA in bone at various temperatures.
(c) Variance in DNA preservation
Only 38.6 per cent (figure 3) of the variation in DNA preservation could be explained by the age of the fossils. Despite efforts to minimize variation (see §2), part of this variance can be ascribed to qPCR stochasticity. Results from qPCR repeats of 50 DNA extractions showed an average of 0.9 cycle shift in CT value (see the electronic supplementary material, table S5). In 36 of the 50 repeats, the two qPCRs deviated with less than one cycle, confirming the general consistency in the set-up. The 50 qPCR repeats spanned a total of 14.8 cycles in CT, implying that roughly 6 per cent (0.9/14.8) of the total variation in CT could be ascribed to qPCR stochasticity. This value is the procedurally introduced background noise in our estimated decay rate. Lastly, with regard to accuracy, it is possible that low-template aDNA extracts (typically with CT > 35) could include an amplification signal of template fragments forming as a result of incomplete extension (and where fragments eventually forming a full length product in latter PCR cycles). Such fragmentary chimaeric templates forming as a qPCR artefact might subtly impact our estimates of decay kinetics on the most poorly preserved bones.
The mean CT value for the Rosslea samples (39.6) was significantly higher than those from BHV (31.7; t-test, unequal variances, td.f. = 8 = 4, p = 0.004) and PV (34.1; t-test, unequal variances, td.f. = 7 = 2.9, p = 0.022). When converted to relative DNA copy numbers, PV fossils contained on average 19.6 per cent of the template molecules present in BHV fossils, and Rosslea contained just 0.3 per cent, corresponding to approximately 330 times fewer DNA templates. A 'spike-in' experiment showed that the poor DNA preservation at Rosslea did not result from PCR inhibition (see the electronic supplementary material, table S2). Whereas poorer DNA preservation at Rosslea may be explained by its greater age (figure 2), the difference between the largely contemporaneous PV and BHV sites is more intriguing. This difference was significant, not only when including data from the entire combined PV-BHV time frame (t-test, equal variances, td.f. = 148 = 4.5, p < 0.001), but also within the 1882 years of temporal overlap (915–2797 BP, n = 72). Eliminating age as a contributing factor, PV fossils still displayed significantly higher CT (i.e. poorer DNA preservation) than those from BHV (t-test, unequal variances, td.f. = 72 = 3.8, p < 0.001).
We tested whether these site-specific differences in DNA preservation could be explained by differing decay rates. After log-transformation, PV (R2 = 0.359, ANOVAd.f. = 102: F = 56.53, p < 0.001) and Rosslea (R2 = 0.749, ANOVAd.f. = 7: F = 17.91, p = 0.005) both displayed highly significant regressions, whereas the same decay signal was not supported for BHV (R2 = 0.058, ANOVAd.f. = 46, F = 2.79, p = 0.10; figure 3). The average molecular half-lives for DNA in bones from PV and Rosslea were 506 years and 389 years, respectively. However, an analysis of covariance (based on standard error of difference between slopes) showed that the decay rate was not significantly higher at Rosslea than in PV (td.f. = 107 = 0.92, p = 0.36) or the whole dataset (td.f. = 162 = 0.70, p = 0.48). It was therefore justifiable to pool the data from the three sites to achieve an average DNA decay rate for DNA in a North Canterbury bone (figure 3). We could not detect an exponential decay process in the BHV material; so the decay rate was not estimated for this site alone. Notably, whereas 74.9 per cent of the variance at Rosslea was explicable by geological age, only 5.8 per cent was explicable by age at BHV (figure 3). The discrepancy probably reflects differences in the periods of deposition at the sites (shorter at BHV), allowing geological age to dominate the relationship at Rosslea but not at BHV.
We then examined the effect of storage time. The difference in average CT between PV and BHV could be an effect of the seasonal water saturation at the PV site, but it could also be a storage effect, given that the material was collected from BHV in 2001, whereas most of the PV material had been at Canterbury Museum for more than 50 years (figure 2). However, we note that the DNA in bones from Rosslea was more degraded, despite the bones having been sampled shortly after excavation. Moreover, the range in CT (figure 2) was highest at BHV although the 47 bones from this site had identical storage times (7 years between excavation and sampling). A previous investigation showed that storage conditions can be a major contributor to variation in DNA preservation [50]. A multiple regression analysis showed that both variables (age and storage) had a significant effect (p < 0.001 for both), but whereas in combination, they explained 45.2 per cent of the variation in DNA preservation, only 7.7 per cent of this could be ascribed solely to storage time. Therefore, in our experimental setup, the geological age of a sample was a far better predictor of its DNA preservation than storage time.
(d) Decay rates from Illumina data: mtDNA versus nuDNA
Illumina sequencing of two DNA extracts yielded 10.1 and 24.8 million reads, respectively. When filtered and mapped against a draft of the ostrich genome (see §2), the return was 3128 and 34 495 unique, non-duplicated reads (see the electronic supplementary material, table S6). These should represent endogenous fragmented moa DNA and hence suitable for calculations of DNA decay rates. It was first confirmed that the number of fragments declined exponentially with fragment lengths (see the electronic supplementary material, table S6 and figure S3), thereby conforming to the random fragmentation model [23]. The DNA decay rate (k) was calculated as 2.13 × 10–5 and 2.71 × 10–5 per site per year, for each sample, respectively (figure 4). The majority of DNA within most vertebrate cells is derived from the nucleus, thus it was a priori hypothesized that the majority of the mapped 'shotgun' reads would represent nuclear DNA fragments. If there is a difference between post-mortem DNA decay rates in the nuclear and mtDNA genomes, it could explain why the rate is four to five times faster than estimated from our qPCR data (5.5 × 10–6 per site per year), which represent fragmentation of mtDNA.
To investigate this further, the shotgun sequences were then mapped against the moa (D. robustus) mitochondrial genome, returning 878 and 4032 reads, respectively. When the decay rate was calculated based only on these mtDNA sequences, it was found to be 2 and 2.5 times slower, respectively, than that estimated from the entire dataset (figure 4, electronic supplementary material, table S6). These numbers suggest that mtDNA degrades at a slower rate than nuDNA, consistent with observations reported in the study of Schwarz et al. [24]. This could perhaps be explained by the circular structure of mtDNA, making it less accessible to exonuclease activity.
The two decay rates of mtDNA measured from Illumina data appear slightly faster than the rate based on qPCR (figure 4). This could be sampling error because these are just two individual data points compared with the qPCR average of 158 samples. However, it could also be explained by the fact that the qPCR decay rate is based on differences in DNA preservation between bones that have all been in the ground for more than 600 years. This rate does not therefore incorporate the presumed faster DNA degradation occurring in the initial post-mortem phase. It has also been demonstrated that the library preparation for high-throughout sequencing may skew the fragment length distribution towards shorter average reads [66], and such bias could potentially explain the observed rate difference between our qPCR and HiSeq data. However, it is not clear whether such bias would affect only the average read length, or whether it could also alter the slope of the frequency distribution, which we use to estimate k. Further data are required to clarify these potential biases.
4. Concluding remarks
We have demonstrated that in situ DNA decay is described by first-order kinetics, confirming that long-term post-mortem DNA fragmentation can be treated as a rate process. This closes the gap to theoretical in vitro observations made four decades earlier. We argue that equation (3.3) represents the best available approximation of the rate of mtDNA decay in fossil bone. Empirical data from a range of different depositional environments, also extending back into the Pleistocene, are now needed to further test and refine this model.
Importantly, we show how high-throughput sequencing data can be used to estimate DNA decay rates, and thus the vast amount of such data currently being generated worldwide may provide an excellent resource by which to study DNA decay. Factors such as bone thickness, burial depth, surrounding pH and water saturation can be tested and modelled in future research, whereas information on season of death, speed of cadaver incorporation into sediment and climatic fluctuations will probably be less accessible.
It is tempting to suggest that we can now predict the temporal limits of DNA survival, and finally refute the claims of authentic DNA from Cretaceous and Miocene specimens. This is, however, not straightforward. One needs information on the number of template molecules in living tissues, and estimates of post-mortem DNA decay rates for each tissue type. However, the half-life predictions (table 1) display the extreme improbability that an authentic 174 bp long mtDNA fragment of an 80–85 Myr old bone could have been amplified [1].
Our results indicate that short fragments of DNA could be present for a very long time; at –5°C, the model predicts a half-life of 158 000 years for a 30 bp mtDNA fragment in bone (table 1). Even rough estimates such as this imply that sequenceable bone DNA fragments may still be present more than 1 Myr after deposition in deep frozen environments. It therefore seems reasonable to suggest that future research may identify authentic DNA that is significantly older than the current record of approximately 450–800 kyr from Greenlandic ice cores [47].
We are grateful to Malene Møhl for help with the sampling and to Emma McLay, Helen Hunt and Jayne Houston for technical assistance, and to Søren Overballe-Petersen for useful discussions. We thank the Hodgen (PV), Giesen (BHV) and Earl (Rosslea) families for their support of our research. Thanks to Oliver Ryder, San Diego Zoo for providing ostrich tissue. Financial support was provided by the Marsden Fund of the Royal Society of New Zealand (contract 06-PAL-001-EEB to Palaecol Research Ltd). M.C. and D.H. received support from the SYNTHESYS project (http://www.synthesys.info/), financed by European Community Research Infrastructure Action under the FP7 'Capacities' programme. M.T.P.G. and J.A.S. were supported by the Danish Council for Independent Research (Grant 10-081390). M.B. is funded by the Australian Research Council (Future Fellowship FT0991741).
- Received July 26, 2012.
- Accepted September 14, 2012.
- This journal is © 2012 The Royal Society


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Ryan, Obama and 'Racism'
Two men use similar language and the left calls one a bigot.
A week later, and liberals are still lining up to assail Paul Ryan's "racism." The episode is worth noting not because Mr. Ryan said anything wrong, but because of what it shows about the political habits of today's elected and media left.
The Wisconsin Congressman has been looking into the problem of upward economic mobility and how effective federal programs are in combatting poverty. Appearing on Bill Bennett's radio program, Mr. Ryan observed that antipoverty assistance can often create "incentives not to...

subset: commoncrawl


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Published on January 28th, 2013 | by behostels0
Barcelona's Carnival 2013 is coming!
From Feburary 7th to the 13th, Barcelona becomes a color filled city when carnival takes over the streets. People from all around the world come to Barcelona to enjoy this unforgetabble party! Are you going to miss it? Be Hostels wants you to enjoy this great event with us in Barcelona!
Carnival lasts a week and is a time of complete joy and exuberance before Ash Wednesday. People dress up in special costumes and disguises and there are parades and events all across the city. In Barcelona, the carnival begins on ´Greasy Thursday´ (´Dijous Gras´). This day is celebrated by eating good and excessive meals. The big Carnival Parade will be held during the weekend. Then, on Ash Wednesday, Carnival comes to an close with the 'Sardine's funeral' when the King of Carnival dies and is buried. Colorful costumes are exchanged for black clothes and tradition calls for eating fish.
So…you can expect some fun days and crazy nights! The carnival boasts a number of colourful parades with scantily clad revellers, outrageous costumes, street music, and bands.
You can also visit the Sitges Carnival, near Barcelona, which has become world-famous. This Costa Dorada destination manages to pull off one of the most amazing Carnaval experineces possible, year after year- an extravaganza of colour and hedonism that can rival any city's celebrations. Sitges is just a 40-minute train ride away from Barcelona, and a great option for a day (or night!) trip out of the city.
The best way to get to Sitges from Barcelona is by train. There is frequent service from Passeig de Gràcia and the Estació de Sants.


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|Music: bring down music.
Guess where I'm at. I am at California State University Long Beach and this pyramid behind me doesn't hold mummies. It actually holds athletes. Pretty unique, Huh? I'm Nicole Erin and lets start this tour.
Campus montage.
California State University Long Beach is located in Long Beach, California.
Valerie Bordeaux
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Voice Over host.
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Lisa Vollendorf
We have really rich, varied discussions because the students all come from different walks of life and share their experiences in a really mutually respectful way.
Valerie Bordeaux
We're ranked among the top ten universities in the nation as far as applications to the university. Our students are highly competitive in terms of the typical GPA. We're talking about a 3.5, about 1,100 in terms of their SAT scores.
Voice Over host
For a full-time, in-state, undergraduate student the average cost of yearly tuition is about $5,500, while those coming from out of state will pay the same plus $372 per unit. Check out the President's Scholar Program, which offers full scholarships including housing costs.
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CSU long beach has consecutively ranked as one of the top 5 public comprehensive universities in the western u.s. by U.S. news and world report. Princeton review agrees naming it one of the best in the west!
Voice Over host
The university offers 86 baccalaureate degrees, 67 masters degrees and 2 doctoral degrees in 7 academic colleges, as well as 74 minors and 38 programs leading to education credentials.
Valerie Bordeaux
We're the largest single campus university in the CSU, but although we're the largest, We do have small learning communities so student don't feel like they're a number when they come onto the campus. As I mentioned they all go through student orientation programs so they have a nice successful transition. They feel comfortable with the university.
Lisa Vollendorf
We have a great commitment also to advising both in our programs but across the university I would say it's a great big university in many ways but it has a very small university feel.
Chris Lowe
Our students get kind of one on one interaction with the faculty. They get to know all the other students with in their program and ofcourse they have access to all these great beach and marine environments only three miles from campus.
They're very good at working closely with you and not only helping you to not only have the knowledge but the necessary skills to survive outside and I' going to grad school so my professor in the marine biology department has met with me for hours just in preparing me helping me get good personal statements and also giving me the skills in order to do research in the future.
Voice Over host
The 5 most popular baccalaureate programs are liberal studies, communications, studio art, nursing and biology.
Chris Lowe
So the great thing about teaching marine biology is that it's a very hands on field. SO we get to take our students out in the field. We do a lot of field trips. They get to go out in boats and many of the thing we are teaching them reinforces what they learn in lecture but it also gives them that kind of hands on experience that they're going to need to get a job in this field.
Voice Over host.
Cal state long beach profs stay connected to their students keeping class sizes small. In total there are 1886 faculty members, about half of whom are full–time.
The professors are just phenomenal. I just can't say enough good things about them. They are willing to meet with you at any time of the day. Their doors are always open and they will help you with whatever's going on whether it be a school issue, a life issue or whatever. They're always there, and they are willing to be not only your professor but your friend.
Go beach! you'll hear that chant as the underclassmen cheer on the 49ers as the 18 varsity sports compete in the big west conference at the div. 1 level.
Give it up for the ladies! The volleyball team has won 3 national titles, 1 national runner–up and 8 final 4 appearances!
The beach has 18 residence hall buildings accommodating over 2,400 students. Living on–campus costs about $10,000 a year depending on what meal plan you purchase.
You can even get academic assistance in the Residence Halls. Residents take advantage of the learning communities, on–site classes, advisement and tutoring services!
Valerie Bordeaux
We have over 300 different clubs and organizations that students can get involved in. We have a fabulous, new state-of-the-art rec and wellness center that just opened for our students.
Notable facts/future
Voice Over host.
Cal state long beach breeds some of today's most elite athletes. MLB sluggers Jason Giambi and Evan Longoria both attended the university.
A natural fit for the beach, professional beach volleyball player and 2 time olympic gold medal winner Misty May–Treanor is an alum.
Funny man Steve Martin has received degree.
Plus, academy award winning director Steven Spielberg has received degree!
Host standup/close/goodbye
it's hard to choose my favorite place on campus, but this Japanese garden is pretty much on top of my list. What a perfect way to end this tour. I'm Nicole Erin for University TV. Beach Pride!
View Youniversity TV's tour of CSULB here.View alternative content


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(Dana White establishing a neural link with the production truck. / Photo via Getty)
We'd say Ronda Rousey was the hammer and Alexis Davis was the nail at UFC 175, but that doesn't even come close to how badly Rousey destroyed her Canadian opponent. Instead, we'll put it this way: Ronda Rousey was a 500 kilaton hydrogen bomb and Alexis Davis the arid desert or frigid Siberian tundra it was tested on.
The fight wasn't a fight. Calling it a squash match doesn't even convey how one-sided it was–that's how one-sided it was.
Since Rousey, at least as far as the world knew at the time, suffered no damage in her 16-second trouncing of Davis, people thought she might be able to save the day at UFC 176, a card desperately in need of a main event.
One of the people who thought this was a UFC production truck employee. During Rousey's post-fight interview, Rogan claimed the truck asked him to inquire as to whether Rousey would want to fight at UFC 176. She was friendly but gave a diplomatic non-answer, stating she needed knee surgery and it depended on what her coaches said.
Dana White, however, was not so friendly about it…
(via r/MMA)
He'd later tell ESPN's Brett Okamoto that Rousey being asked that question was the "biggest F up ever."
Damn. We know White has a flair for the dramatic, but this is a bit over the top. Biggest F up EVER?
What ever you say, Uncle Dana!


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Milton Rogovin: Social Documentary Photographs
Artstor is collaborating with The Rogovin Collection to share 260 images of Milton Rogovin's social documentary photography in the Digital Library. A documentary photographer and political activist, Milton Rogovin (b. 1909) is best known for his portraits of the poor and working class, often considered "the forgotten ones," and his depictions of their lives, communities, and working conditions. Trained as an optometrist, Rogovin turned to photography when his political activism drew the attention of the House Committee on Un-American Activities in 1957 and negatively impacted his practice in Buffalo, NY. In 1958, Rogovin began his first photographic series, documenting Store Front Churches in the African-American community in Buffalo's East Side. He continued to photograph the working people and ethnic communities in the Buffalo area, often photographing his subjects both at their workplaces and in their homes. Starting in 1972, Rogovin photographed families in the area near his optometry office in Buffalo's Lower West Side, the city's poorest and ethnically-diverse neighborhood. Over the course of three decades, Rogovin would return to his subjects, in some cases tracing the same family's history in a series of triptychs and quartets. Further afield, Rogovin explored the plight of workers, particularly miners, in the small towns of Appalachia. In 1983, Rogovin received the W. Eugene Smith Grant in Humanistic Photography, which allowed him to expand his "Family of Miners" series to include workers in Scotland, France, Spain, Germany, Czechoslovakia, Cuba, Mexico, China, and Zimbabwe.
Throughout his career, Milton Rogovin's work has appeared in more than 160 journals, magazines, and other publications. Rogovin has participated in 60 solo exhibitions and more than 30 group exhibitions. His prints and negatives have been acquired by the Library of Congress and the J. Paul Getty Museum. His master set of photographs is at the Center for Creative Photography at the University of Arizona, Tucson.


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Education and the economy are among the top issues expressed by the three candidates seeking to serve District 59 in the Indiana House of Representatives.
Appearing on the Nov. 6 general election ballot are Republican Ryan Lauer, Democrat Dale Nowlin and Libertarian Clyde Myers, all of Columbus. The winner will fill the seat held by Milo Smith, a six-term lawmaker from Columbus who chose not to run for re-election.
District 59 encompasses Columbus, Taylorsville and in general areas west of State Road 7 in western and southern Bartholomew County.
Lauer, who ran against Smith and lost in the 2014 and 2016 Republican primaries, said it would be his highest honor to serve the people of Bartholomew County in public office.
"I believe strongly in service to our community and making a positive difference for the lives of all Hoosiers," Lauer said. "I will serve earnestly, honestly and with integrity seeking, seeking what is right, listening to the people and protecting our Constitutional rights."
Nowlin, a math teacher at Columbus North High School since 1985, said as he became involved with issues in downtown Columbus, he saw that his efforts could have a positive impact on his community.
This is his second attempt to win the District 59 seat. Nowlin mounted his first campaign for the seat in 2016, losing to Democrat Bob Pitman in the primary.
"As an educator, I knew we could do more to support and improve our schools," Nowlin said. "I decided to use my people skills, my understanding of data and my expertise in education to serve our wider community."
Myers said he was motivated to run for office since he contends that Indiana lacks no champions for businesses.
"Republicans and Democrats continue to move the goalposts on businesses and individuals trying to work in our state," he said. "They pass legislation without any analysis for unintended consequences. I want to legislate using facts, data, reason, logic and principle, not emotion, opinion or party bias."
The Alliance for a Healthier Indiana is lobbying to increase the statewide tax on cigarettes — currently 99.5 cents a pack, set in 2007 — by $1.50 a pack in hopes of reducing smoking among Hoosiers and trimming long-term healthcare costs.
However, all three candidates said they are opposed to increasing the cigarette excise tax at that level.
Nowlin said he doesn't support an increase of that magnitude, noting that Indiana has one of the highest smoking rates in the nation.Nickels Womens Light Gray/ Off White Leather Loafers Size 9 MSperry Top Sider Women Tan Leather Boat Shoes Sz: 7.5 , BELINDA New Fashion Blink Slip On Casual Women Flats Beauty Bowknot Shoes Silver , Munro American Black Strand Fabric Leather Slip On Shoes 1.5" Block Heel Sz 10m , Women batman Flats Shoes DC Comics Slip On Sz M Canvas Casual Licensed NWT$79 NEW Nine West RUMBA Slip On Suede Sneakers Turquoise Black Shoes 8Dr Scholl's Reply Slip On Shoe - Women's Size 7.5M, BlackZara authentic genuine ponyhair fur leather shoes women's 6 36 slip onClarks Women's Valley Moon Flat,Fuschia,9.5 M US , Women's Nike Lunarsolo Running Shoe, Rose Gold, Size 9 , Saucony Cohesion 7 Running Athletic Shoes Hot Pink Trim Size 6-1/2 15191-18 WideKeds Women's Champion Wash Stripe Ankle-High Canvas Flat Shoe , Stuart Weitzman Womens 8.5 M black patent leather ballet flat shoes pump slip-onConverse Chuck Taylor - Mint Green - Men's 7 / Women's 9 , New Balance 758 Silver & Purple Pink Athletic running Shoes Womens Sz 9 , Reebok Womens Renaissance White/Steel Fashion Shoes Size 8 (359359)BLOW OUT SALE @ US POLO ASSO Hi Tops ZEBRA NEON Sneakers Womens Shoes Sz 11 ??b4New Balance 880v3 Womans Running Shoes- Size US9- Blue/Orange RRP$219 , Nike Womens Nike Zoom Strike Running Shoes Size 6.5 (15595) , Brooks PR MD 54.26 Women's Middle Distance Track Shoes 1200871B395 MSRP $65AGC-95A Asics Gel Contend 2 Women's Athletic Shoes, Size 9.5 M USWOW NEW 8.5 Converse Chuck Taylor All Star Animal Leopard Cheetah Safari PrintWomen's NEW CULT WOMENS SNEAKERS Discount Athletic ShoesNew Balance Women's 670v5 Shoes Pink with NavyWomen's Brooks Launch 4 1202341B077 Gray/Turquoise Running Shoes Size 7.5 MConverse CTAS Madison OX 560637C Ocean Bliss Aegean Storm Casual Shoes WomenNIKE Ladies MERCURIAL SOCCER CLEATS US 9.5 , Onesole Women's Patriotic Double Combo Multicolored , New Skechers Womens Street Freestyle Platform Black Leather Shoe Sneaker sz 8MSkechers Blue & Green Mesh Bungee Slip-on Sneakers Glider Twilight 7W New
Nearly 25 percent of Hoosier adults smoke, compared with 17 percent nationally.
"Indiana's cigarette tax would be higher than any surrounding state, resulting in other economic consequences," Nowlin said. "I would not support any increase unless all new revenue went towards smoking prevention and support for those trying to quit smoking."
Lauer cited information from the Centers from the Disease Control and Prevention that indicated most smokers are under or near the poverty level.
"Raising this tax will have a disproportionately negative effect on lower-income Hoosiers," Lauer said. "Attempting to change people's behaviors and addictions by raising taxes is ill-conceived and will hurt our citizens with lower incomes."
Myers agreed, referring to the cigarette tax as nothing more than a tax on poor people.
"It's insulting to people who are already struggling to kick them down. No, I do not support a cigarette tax nor excise taxes of any kind," Myers said.
The three candidates also offered their perspectives on different topics.
Q: Identify three priorities for the next legislative session and why have you chosen them?
Lauer: "Attack the opioid and methamphetamine epidemic by increasing enforcement, awareness and treatment options. We are all affected by this scourge and too many children and families have suffered. Reform the Department of Child Services. Serious issues have been identified at DCS failing our most vulnerable children who must be protected. Expand school choice with education savings accounts for all children and allow more local control of education by letting teachers teach again."
Nowlin: "Indiana can be a welcoming state with a thriving, sustainable economy. That requires a well-educated workforce. We can grow that workforce by investing in Hoosiers and by creating a quality of life that will attract newcomers. That means we must fix the Indiana Department of Child Services, support and improve our education system and support local communities as they work to overcome the opioid epidemic."
Myers: "Business — The Indiana General Assembly consistently props up some businesses and industries, while simultaneously putting roadblocks in front of others. If we want good jobs in this state, we need to legalize work and let people earn their own living without a mountain of bureaucracy in their way. Education — Many seem to think that the more money spent on education, the better. We may indeed need more money for education, but that isn't the only factor. There are other states performing significantly better without many more resources. We need to implement what works, not just throw money at problems. Personal liberty — I am the only candidate running that doesn't think that he knows better than you how you should run your life. I think you own yourself, your body, your property, your business, your relationships, your self-defense. No one else should decide how you should live, so long as you aren't hurting anyone else."
Q: What is your stance on Indiana enacting a hate crime law and explain why you feel that way?
Lauer: "Judges today under current law have wide latitude to increase penalties for crimes with hateful motives and should do so. I support harsher penalties for violent crimes and heroin and meth dealers who are killing people. All violence is hateful and motive should be considered in sentencing, but hate-crimes proposals do not cover all biases and hate."
Nowlin: "I believe Indiana should pass a hate crime law. It would send a message that all people are welcomed and valued. That is a message for members of minority groups in Indiana and people outside of Indiana, who might be considering making Indiana their home. It is the right thing to do and it would provide more quality candidates for Hoosier companies."
Myers: "I wish things were so simple that we could just legislate away hatred and bigotry. I would be in favor of legislation that has undergone a thorough analysis to protect against it backfiring and hurting those it is designed to protect and does not infringe on constitutionally protected freedoms."
Name: Ryan Lauer
Age: 41
Current office held: None
Previous offices held: Bartholomew County Council, elected in 2010, president in 2014
Education: Bachelor of Science in biochemistry, Indiana University
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Community: President of Grace Lutheran Church Council and Congregation, Grace Lutheran church member, sheriff's merit board member, Community Corrections advisory board member, co-founder of "Save Eos," co-chair of Uncommon Cause 2012, Columbus Philharmonic violinist, computer and data processing board member, Minds on Math mentor at CSA-Lincoln, Columbus Arts District retail committee
Family: Wife Blair, daughters Awyn and Lillian and son Isaac
Name: Dale Nowlin
Age: 64
Current office held: None
Previous offices held: None
Education: Bachelor's degree in mathematics and psychology from Alma College, master's degree in mathematics from Michigan State University
Community: Teacher and math department chair at Columbus North High School and Northside Middle School, sponsor of the Columbus North Environmental Club, Historic Downtown Neighborhood Alliance Steering Committee, member of the Downtown Working Group, First Presbyterian Church Elder and chair of the Adult Fellowship and Education Committee, Ivy Tech Columbus Advisory Board member, former president of the Indiana Council of Teachers of Mathematics, former volunteer for Campus Life/Youth for Christ
Family: Wife Isabel, three grown children, Elisabeth, Joel and Matthew.
Name: Clyde Myers
Age: 41Dolce Vita Shoes 6 Brown Black Calf Flats Hair Hide Zebra Print Flats Oxfords Shoes Sz 6 48f858b
Current office held: None
Previous offices held: None
Education: Associate degree in computer networking from Ivy Tech, 1995 graduate of Columbus North High SchoolFlats
Family: Wife Amy, daughters Margaret and Olivia
Meet the candidates for Indiana House District 69: Republican incumbent Jim Lucas and Democrat Steve Schoettmer.


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|The authors own nothing. Joss, UPN, WB, etc. own Buffy, the show, the characters, the places, and the backstory. The authors own any original plots.


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» Tires » Hankook » Optimo H426
Price start from $86.61 + Tax
Passenger Cars / All Season /
Designed to provide comfort in driving and elegant style. Optimo® H426 employs 5-pitch design to optimize lateral grooves to drastically reduce noise. Rounded main groove edge minimize noise arising from the vibration of block edges.
- Optomo® H426 uses an advanced tread silica compound that improves both wet braking and fuel economy.
- New hydrophilic material features superior wet grip and braking performance compared to previous carbon compounds .
- Test results show that Optimo® H426 provides lower rolling resistance and better wet and snow performance.


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DistributionRead full entry
Australian sea lions are found on islands offshore of Australia. They range from western Australia to islands in southern Australia. The largest populations are found on Kangaroo Island and Dangerous Reef (near Port Lincoln) in southern Australia. The smallest numbers are found on the west coast of southern Australia and in Western Australia. Some live in Tasmania as well, but these are few in number. They once bred in the Bass Strait, but the entire population was wiped out by the sealing industry.
Biogeographic Regions: australian (Native ); indian ocean (Native ); pacific ocean (Native )


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OK- I have a pretty good buzz on. [and I wanna cigarette- damn nasty habit. Aint smokin none tho.] Dice- I don't have any more I-Ron City Aluminum bottles left. Thats gooda beer. [kinda gassy, I burped like.... well alot] The IC I won in cans I didn't like but this was real smooth beer. Thank you-
Jerry- I bought 2 six packs so I could send MaryLou some "Sunshine Wheat" with your "Fat Tire" and the box only held 8 or 9 of them [I cant member] So we drank the resat of those too. Mmmmmmmmm your getting some good beer!


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Thank you, Tom Menino, for taking the time to wade in on Elizabeth Warren's real or imagined Native American heritage, for letting us know that we need not be concerned about it because there are more important issues to confront, things like crime, education, and jobs.
It sounded so adult, so mature, that I wanted to stand up and shout, "Hear, hear! Finally, a voice of sanity in the asylum.'' But there's one small problem with Menino's point, and that problem is this: Our normally sage mayor is wrong.
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FOREWORD v
THE Two GREAT GODS OF EARTH
How THE WORLD AND MANKIND WERE
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Achilles Admetus Aeneas Agamemnon Aphrodite Apollo Apollodorus aqj puB aqj ui Artemis asked Athena bade beautiful blood brother called Clytemnestra creature cried dark daughter dead death Dionysus divine earth Euripides evil father fear fell Fleece forever gave give goddess gods golden Greece Greek hand heart heaven Hector Hera Hercules Hermes hero Hesiod horses husband Iliad Iphigenia Jason jBqj killed King knew land lived looked maiden married Medea Menelaus mortal mother mythology never night nymph Odysseus Oedipus Olympus once oracle Orestes Ovid palace Pelias Pentheus Perseus poets Poseidon Prometheus Psyche punished qjiM sail SBM aq seemed ship sister snaz spoke spoS story Telemachus tell terrible Thebes Theseus things thought told took Trojans Troy turned uiiq uiojj Venus wife wind woman women words young Zeus Zeus's


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Third child of Princess Juliana, Princess Margriet is born in Ottawa, Ontario. View of Netherlands flag flying on Parliament Hill. Prince Bernhard carrying his daughter walks out of a room in a hospital at Ottawa. Princess Margriet's name on the registration document is shown. Shows Prince Bernhard with his daughters in snow.
President Eisenhower visits Ottawa, Canada. President of the United States Dwight D. Eisenhower and his wife Mamie Geneva Doud Eisenhower get off an aircraft at Ottawa. Officers greet and receive them. He arrives in Ottawa to talk to Prime Minister of Canada John George Diefenbaker and Canadian leaders. Eisenhower addresses the parliament on difference in trade policy. Gathering applauds.
U.S. Air Force Chief General Hoyt S. Vandenberg in Ottawa, Canada. U.S. Air Force C-121 aircraft taxis along the airfield. USAF Chief General Hoyt Vandenberg talks to an officer at an airfield in Ottawa. The General greets the dignitaries who arrive for the Ottawa conference.
King George VI and Queen Elizabeth of the United Kingdom in Ottawa, Ontario, Canada. A large crowd gathers on a street to greet the King and the Queen. Bakers standing near a table. A cake on the table. The King and the Queen come outside a building. A large crowd cheer and wave towards them. People standing near the cake. Aerial view of the large crowd on the street. A war memorial at Connaught Square dedicated to the soldiers who died in World War II. The dignitaries place a wreath on the memorial.
Alexander Cambridge arrives as the new Governor General of Canada in Ottawa, Ontario. Officers greet Governor General Alexander Cambridge. An aircraft flies over a clock tower. Honor guards lined up outside the Parliament Building. Governor General with officials. Governor General with his wife Princes Alice as she holds flowers in her hands.
Brief unrelated view of a man plastering a wall to repair it. Then: U.S. Air Force Chief General Hoyt S. Vandenberg in Ottawa, Canada. USAF Chief General Hoyt S. Vandenberg talks to various officers on the tarmac of an airfield in Ottawa. The General greets the dignitaries who arrive for the Ottawa Conference. Both men and women are seen . General Vandenberg talks to an officer as he smokes a cigarette. An aircraft in the background. Women talk at the airfield. Officers enter a building at the airfield.


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Leonardo DiCaprio: TAG Heuer Event with Cameron Diaz!
It's a Gangs of New York reunion!
Leonardo DiCaprio makes his first appearance since his split with girlfriend Erin Heatherton at an event for TAG Heuer on Saturday evening (November 10) in New York City.
The 37-year-old actor helped his former co-star Cameron Diaz celebrate the launch of her limited edition signed watch at the event which raised money for the UN Women and New York Cares' Hurricane Sandy relief effort.
Leo's split from Erin was announced last week, just days before she walked in the Victoria's Secret Fashion Show.
FYI: Cameron is wearing SLANE bangle bracelets.


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Looking for work in Australia? Well look no further -- here are the Top 10 great places to work down under, based on BRW magazine's list which used research conducted by the Great Place to Work Institute which surveyed more than 15,000 employees.
The magazine said there was a common theme at all the Top 50 companies - perks. Benefits vary from free iPods, Krispy Kreme doughnuts and the opportunity for international travel.
Popular firms also offer such benefits as education subsidies, on-site fitness centres or subsidized gym memberships, flexible work arrangements including telecommuting, health insurance plans, senior management opportunities for women and child care.
By Dora Cheok and Zara Fung
Posted 18 May 2009


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ince 1984, our mission has been to provide a wide range of certified organic grocery products to retailers and distributors throughout North America. We are dedicated to the development and growth of a worldwide network of organic growers, suppliers and processors based on the concepts of organic, sustainable agriculture.
Our collection of branded organic grocery items includes some of the leading sellers in their respective categories. Villa Flor Extra Virgin Olive Oil, Montebello Artisan Italian Pasta, Lorenzi Balsamic Vinegar from Modena, Nunez de Prado Extra Virgin Olive Oil from Spain and Lapas Extra Virgin Olive Oil & Olives from Greece all enjoy prominent positions on the retail shelf because of their quality and packaging. See Branded Products for more information.
We provide a full range of turnkey private label products to retailers and distributors nationwide. Our vast knowledge of organic markets allows us to offer the most competitive pricing and unparalleled service to our private label customers. See Private Label for more information.
Spruce Foods


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Welcome in the Website of ECP Italy
ECP – Europe for Citizens Point Italy is the National Contact Point for the "Europe for Citizens" 2007-2013 Programme.
By Decision 1904/2006/EC of 12 December 2006 the European Parliament and the Council adopted the "Europe for Citizens" Programme, that covers period from 2007 to 2013 and that has an overall budget of 215 million euro.
The Programme promotes – through financial supports – projects and organisations that foster active European citizenship and therefore the involvement of citizens and civil society organisations in the process of European integration.
Download the publication "Fostering the Dialogue between Citizens, Civil Society Organisations, National and European Institutions. An Introduction to the European Year of Voluntary Activities promoting Active Citizenship European" (PDF, 3357 kb)
Busseto: Current and future skills in the health and welfare sector
19 April, 2012
Roma: Fundamental Rights, European Remembrance and Holocaust. An introduction to Action 4 of the Europe for Citizens Programme
16 April, 2012
Firenze: Info day Programma Europa per i Cittadini 2007 - 2013 e Opportunità per gli enti locali
23 April, 2012


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text: After the Indians filled their holes in the bullpen and in the infield (for the short and long term) with a sizzle, it seems that the off-season will end with a bit of a fizzle as the Tribe has signed Carl Pavano to the 2009 rotation mix on a one-year, incentive-laden deal that guarantees Pavano $1.5M with $5.3M additionally available if certain incentives are reached, ostensibly ending the Indians' off-season. The rationale behind the Pavano deal is that adding him to the rotation gives them some depth in the rotation, allowing Pavano to be a candidate to fill one of the three spots in the rotation behind Lee and Carmona. Since a number of the candidates for those 3 spots still have options (notably Laffey, Sowers, Huff, and Lewis), it would stand to reason that Pavano (who was dubbed "American Idle" by the brutal New York tabloids), even if remotely healthy or effective, will given a spot out of Spring Training to sit in the #3 or #4 hole in the rotation. A low risk signing with potential to pull that "Kevin Millwood Miracle" of 2005 out of the hat again, right? I guess, but this move doesn't really do much for me as Pavano has really only had two good years (OK, one very good and one decent) in his 11 MLB seasons. While both of those years were his final two in Florida (during which he complied a cumulative line of 3.61 ERA, 1.21 WHIP, 5.78 K/9, 2.08 BB/9, 2.76 K/BB, which is certainly an impressive performance), that was in 2003 and 2004. He's only pitched 145 2/3 innings since then over the last 4 years, and has pitched more than 140 innings only twice (those two final years in Florida) in his 11-year career. When healthy, Pavano's a good middle-of-the-rotation starter...except that he hasn't been healthy in four full seasons now. I suppose if Pavano somehow finds a way to stay healthy, he's another arm to throw into the mix, but his performance (when allegedly healthy) in 2008 doesn't exactly scream that he's turned a corner as a pitcher or that he'll re-discover his 2004 form. He's never been a big strikeout pitcher or had dominant stuff, so he's a decent middle-of-the rotation option...if healthy (which, if I have failed to mention it, is no certainty). It's true that the Indians are buying low on him, like Millwood, and giving him the opportunity to pitch his way into a bigger contract and back into legitimacy, right? Maybe, but prior to Millwood signing his reclamation contract with the Indians (for $7M guaranteed, if you'll remember), he had put up 200+ innings 4 out of his 7 full years in MLB (the lowest inning total being 121) and had put up inordinately better numbers year after year than Pavano, who seems to still be living off the 2004 (which looks to be the aberration when looking at Pavano's body of work) that netted him his payday in the Bronx. Maybe Pavano, out of the bright lights of the big city, can re-capture some semblance of the career that derailed in 2004. I'm not that optimistic that he can, given his varied and illustrious injury history and the fact that his career and reputation have been built on one admittedly fantastic season. Shapiro's saying all of the right things about his health and his hunger, but the Indians are protected against him being injured or being ineffective enough to be cut loose as the incentives in the contract are pretty straightforward as "Pavano gets $100,000 each for reaching 18, 20 and 22 starts, $200,000 each for reaching 24, 26 and 28 starts, $250,000 for 30 starts, $300,000 for 32 starts, $350,000 each for 33 and 34 starts and $400,000 for 35 starts. He gets $100,000 each for reaching 130, 140 and 150 innings pitched, $150,000 each for 160 and 170 innings, $200,000 for 180, $250,000 for 190, $250,000 for 200, $300,000 for 215, $400,000 for 225 and $500,000 for 235." He's had 24 starts twice in his 11-year career and has reached 140 innings only twice, so he's going to have to come ROARING back into shape for most of those incentives to kick in. Being very optimistic, let's say he has 23 starts and 130 IP - his salary bumps from $1.5M guaranteed goes to $1.9M guaranteed. Overall, the signing itself and adding Pavano doesn't do much for me other than to know that there's another arm to add to the rotation mix to perhaps eat some innings until Jake Westbrook returns from injury (hopefully) in the middle of the season. Realistically, a healthy Pavano, is better than simply giving innings to Jeremy Sowers and it adds the depth that the Indians love, where they can go 7 or 8 starters deep, with the top of the AAA rotation serving as insurance against injury or ineffectiveness. It certainly sets up an interesting battle for the 5th spot in the rotation (remember, Reyes is out of options) as Laffey, Sowers, Huff, Zach Jackson (who does have an option left) and Scott Lewis all fight NOT to go to Columbus. But (while I never wanted to remember or invoke this name ever again) let's all say right now that the "Lesson of Jason Johnson" should be in full effect with Pavano. That is, if Pavano is not healthy, or is obviously ineffective, while the talented youngsters that now figure to start the season in AAA (notably, Dave Huff) prove that they're further along than AAA (and Huff may have already proved that in his 16 starts in Buffalo last year), he should be on an awfully short leash and that this low-risk contract should be one that the Indians are not afraid to eat early and admit a mistake before it's...I don't know mid-June or so. More interesting to me than simply adding Pavano is what the signing means in the long-term as it's likely that this is the last move that the Indians make this off-season as they've now added to their bullpen, their infield, and their rotation. Unless Pavano was added to create more depth to allow one of the young LHP to be part of a package for a surer thing in the rotation, this looks to be the 2009 Indians, which brings some things to light. First off, for the first time in what seems like a long time, the rotation figures to go into Spring Training with a lot of "ifs" around each of the principals that figure to make up the rotation: What if Cliff Lee shows that 2008 IS NOT who he is as a pitcher? What if Fausto Carmona shows that 2008 IS who he is as a pitcher? What if Carl Pavano can't stay healthy or has had injuries take their toll on his effectiveness? What if the injuries that shut down Anthony Reyes and Aaron Laffey rear their ugly head again in 2009? What if the Aaron Laffey we saw when he was promoted is nothing more than a Sowersesque mirage? What if Jeremy Sowers' career descent cannot find a bottom? What if the young pitchers like Huff and Lewis fail to take that next step? For an organization that has been designed to be built on strong starting pitching, that's a lot of variables at play to fill out the top 5 spots in the rotation. Perhaps we've been spoiled by knowing who would be getting the majority of the starts during the season when the team got to Spring Training as far back as 2005, but the middle-to-back of the rotation still looks to be built on sand to me, Pavano or no Pavano. Going further than that, the Pavano signing removes, for the most part, the thought of trading Kelly Shoppach for a middle-of-the-rotation starter as Pavano is designed to fill that need. What that means to me is that concerns about the health of Hafner and Martinez and the long-term production of Garko are significant enough that the Indians don't want to part with their insurance policy against another lost or poor year by any of them. Because, on the surface, everyday AB don't look to be there for Shoppach (despite him earning them in 2008) as the Indians' oft-stated stance is that Victor is the catcher, which would mean that Garko remains the de facto 1B. If Hafner is supposed to be healthy for 2009, how does this not suddenly look like a part-time position for Shoppach or maybe some sort of platoon with Garko, when Shoppach far outperformed Garko in 2008? The Indians claim that they can find regular AB for all four players at three positions, but I'm just not seeing how that's going to happen with one of these players being out of the lineup every game. How they're going to balance it out remains to be seen (if, in fact, no more moves are coming) and it will be interesting to see how Shoppach performs in 2009, whether this off-season will come to represent his peak value or if 2008 was simply an appetizer for the main course that Show Pack has in store in 2009. As an aside, this essentially means is that I can stop writing the piece that I was working on suggesting why Ricky Nolasco (FLA), Wandy Rodriguez (HOU), and Mike Pelfrey (NYM) would all be attractive trade candidates for the Indians to target in that their team would be in need of a catcher and each of them represented tangible upgrades (a few years away from FA) over the in-house back-end-of-the-rotation candidates. Regardless, the Pavano signing looks to be it this off-season...so without further ado ladies and gentleman, YOUR 2009 Cleveland Indians!


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Foo Fighters with producer Butch Vig
Photo: John Shearer/WireImage.com
Exploring The Rock Winners
You've seen the list of winners, now take a closer look at the artists winning in the Rock Field categories for the 54th Annual GRAMMY Awards.
Aside from a shared penchant for musical excellence, alt-rockers Foo Fighters and neo-soul diva Adele wouldn't seem to have anything in common. But at Sunday's 54th Annual GRAMMY Awards, the two acts found some common ground as they emerged as two of the biggest winners. By capturing every rock award they were nominated for, Foo Fighters brought their total career tally to 11 GRAMMYs and became the first act ever to sweep the field. When you consider that the award for Best Long Form Video went to director James Moll for Foo Fighters: Back And Forth, the band's showing Sunday seems all the more impressive. Here, we take a closer look at the Foo Fighters' domineering 54th GRAMMY sweep.
Best Rock Performance
Foo Fighters, "Walk"
It was a race between the Foo Fighters, Mumford & Sons, the Decemberists, Coldplay, and Radiohead, but the GRAMMY in this singles-oriented category ultimately went to Foos for their curiously introspective rocker, "Walk." Putting the perfect cap on their GRAMMY win, the band performed a blistering interpretation of "Walk" on an outdoor stage during Sunday night's GRAMMY telecast, complete with a wild-eyed introduction by comedian/actor Jack Black.
Best Hard Rock/Metal Performance
Foo Fighters, "White Limo"
Going up against bands with apocalyptic names such as Megadeth, Mastodon and Dream Theater, and punk rockers Sum 41, Foo Fighters emerged victorious for their punk-inflected track, "White Limo." Similar to many of their songs, "White Limo" finds Grohl and company juxtaposing humor with musical intensity.
Best Rock Song
Foo Fighters, "Walk"
Most rock fans know that Foo Fighters' frontman Dave Grohl initially earned fame as the drummer for grunge rock pioneers Nirvana. But in that band's tragic aftermath, perhaps no one could have guessed that Grohl would go on to front his own post-grunge posse. on Sunday night's GRAMMY telecast, Grohl and company nabbed Best Rock Song honors for "Walk," a track that could easily be interpreted as a nostalgic ode to Grohl's old band. Lyrics such as "Do you remember the days/We built these paper mountains/Then sat and watched them burn" could certainly apply to Nirvana and their penchant for burning the candle at both ends. Additionally, "Walk" was produced by Butch Vig, who also oversaw the recording of Nirvana's magnum opus, Nevermind.
Best Rock Album
Foo Fighters, Wasting Light
The Best Rock Album competition was a tight race, featuring well-deserved nominations for Jeff Beck, Red Hot Chili Peppers, Wilco, and Kings Of Leon. But Foo Fighters took the trophy for their album, Wasting Light. Backstage at Sunday's GRAMMY Awards ceremony, Grohl attributed the album's rough-hewn appeal to the fact that is was recorded in his garage.
"To me this award means a lot because it shows that the human element of making music is what's most important," said Grohl upon accepting the award for Best Rock Performance. "Singing into a microphone, learning to play an instrument and learning to do your craft — that's the most important thing for people to do. It's not about being perfect. It's not about sounding absolutely correct. It's not about what goes on in a computer. It's about what goes on [in your heart] and what goes on [in your head]. Long live rock and roll."


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Residents in Victoria's fire-stricken east are taking in the news that a 75-year-old man has been charged with sparking the massive blaze that killed one person and destroyed at least 21 homes.
Victoria Police said arson and explosives squad detectives charged the Aberfeldy man with five offences, including recklessly causing a bushfire and causing a fire to be lit without written authority in a fire protected area.
Bruce Taylor, whose family defended their caravan park in the Gippsland town of Glenmaggie as the firestorm raged around them, is baffled by the latest allegations.
"I don't know what the mentality is of these people, I'm at a loss really," Mr Taylor told AAP on Wednesday.
"I think what they need to do is get them out on the front line and see if they think they can handle it then - I reckon they might have a different story to tell."
Police said the accused man is to face the Latrobe Valley Magistrates Court on Wednesday afternoon.
The fire started on a rural property in Aberfeldy in Gippsland on the morning of January 17, police allege.
It burnt almost 75,000 hectares of land in Aberfeldy, Heyfield and Seaton, about 200km east of Melbourne.
Stan Hayhurst, 84, was found dead in his car at Seaton on January 18 after the bushfire roared through the town.
A Heyfield resident, who preferred to withhold his name, said whoever sparked the blaze must be "sick in the head".
"Four of my friends have lost houses, and to think one person could start it all," he said.
"How many lives could you lose by starting some silly little fire that can spread more than 200 kilometres?"


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Being overshadowed by Marshall is understandable enough, but I doubt people would really know without looking it up that Thomas was actually drafted ahead of both Bryant and Tebow back in April.
When the Broncos traded back multiple times in April's draft, I thought for sure they would snag Oklahoma State product Dez Bryant with the 23rd pick. After all, head coach Josh McDaniels had nothing but good things to say about Bryant in a pre-draft interview, and it appeared as though he would be a steal with the pick the Broncos had.
When they jumped in front of the Ravens and traded picks with the Patriots, I knew something was up.
When I read the Tweet saying the pick was Demaryius Thomas, I was stunned, to put it lightly. Why would the Broncos take Thomas over Bryant at this point in the draft? Bryant would have been an absolute bargain, and Thomas had been injured leading all the way up to the draft.
I went back to my draft evaluations, pondering the pick.
I looked at scouting reports of the two players, and tried to determine why the Broncos had taken the lesser known Thomas over the big name. It became very clear why Denver made Thomas the top receiver selected in the 2010 NFL Draft, and he's turned into one of the best kept secrets in the NFL.
Kevin C. Cox/Getty Images
At Georgia Tech, Thomas had a career year in his junior season, catching 46 passes for 1,154 yards and eight touchdowns. For those not in the know, that's an average of over 25 yards per reception. What makes Thomas' production last year at Georgia Tech even more amazing?
The Yellow Jackets run the triple option as their offensive set.
It is heavily reliant upon having a running quarterback as well as a powerful running back. Wide receivers are most certainly not supposed to be productive in this type of offense.
As we saw with Calvin Johnson, Demaryius Thomas is simply a special player, and he made the most of his opportunities. Despite Tech averaging over 300 yards rushing per game, he was still able to put together a 1,000 yard season at the wide receiver position, and that is nothing short of stunning.
Still, there were questions surrounding Thomas. Would his foot be able to stay healthy? Would he be able to learn how to run NFL routes?
Some assets that Thomas possesses, coaches simply cannot teach at this level.
The first asset Thomas has that not many receivers have coming out of college is his fantastic blocking ability. Coming out of the triple option offense, Thomas was asked to block a lot, and he's well versed in that regard already, and that's one less thing coaches have to worry about.
Jeff Zelevansky/Getty Images
Instead of coaching blocking, the Broncos can focus on making him a better receiver.
Another asset is his intelligence, and I'll tag humility along with it. To say Thomas grew up in a bad situation is putting it lightly. At the age of 13, Thomas' grandmother was sentenced to 40 years in jail for drug dealing, and his mother was sentenced 20 for being involved with it financially. Demaryius moved around with some of his relatives, first with his grandmother's youngest daughter. He finally settled in with his grandmother's oldest daughter and her husband who is a preacher.
Thomas said when he made the final move, he decided to start living his life the right way, and said that he became a Christian and vowed to always stick to his word. He said one of the things he told his mother was that someday, he would go pro in some professional sport, whether it was basketball or football. That promise was fulfilled when the Broncos selected Thomas with the 22nd pick in the 2010 NFL Draft.
I say Thomas has intelligence, and that is backed up with facts. Thomas had the 10th highest wonderlic score of any player at the NFL scouting combine with a score of 34, which is well above the average for a wide receiver. His humility is very easily detected in his demeanor. He is a quiet player who lets his play do the talking.
Another thing coaches can't teach? Speed.
Thomas reportedly was running a 40 yard dash in the 4.36-4.38 range prior to his foot injury. At 6'3" 229 pounds, the only other receivers in the NFL who might possess that kind of size/speed ratio are Andre Johnson, Larry Fitzgerald, and Calvin Johnson. It's almost unheard of.
Certainly, Thomas is an interesting athletic specimen, but he's having trouble staying on the field. He missed quite a bit of the Broncos' OTA's this offseason, and when he finally got to camp, he re-injured his foot at the Invesco Field practice (on a touchdown catch, no less).
When Thomas finally came back from that minor tweak, he tweaked his foot again in practice and was held out not just the entire preseason, but he was inactive for the Broncos' week one matchup against the Jacksonville Jaguars.
Thomas is again getting healthy, and hopefully the Broncos have allowed enough time this time around for his foot to fully heal. He has been a full participant in practice this week, but he admitted earlier on that he still might be a couple of weeks away. Head coach Josh McDaniels was more optimistic about Thomas' return, giving hope that the rookie could possibly suit up for his first game this weekend when the Broncos host the Seattle Seahawks.
Whatever the outcome, the Broncos simply need to wait until Thomas is at 100 percent.
He is going to be an asset to this team when he is fully healthy, and outside of his broken foot in the offseason, Thomas had absolutely no trouble staying healthy at Georgia Tech and is not by any means an injury prone receiver, and hopefully it doesn't become that way either.
What the Broncos lost in Brandon Marshall, they gained back and then some in the talent of Demaryius Thomas. He was absolutely dominant for the most part at camp, and caught two touchdown passes in scrimmage action at the Invesco Field practice before he re-aggravated the foot injury.
Still, nobody in the NFL seems to be talking about him.
That's good news for the Broncos, who would just as soon keep Thomas one of the NFL's best kept secrets and unleash him at the expense of an unexpecting opponent.
Thomas was the first receiver selected for a reason. This week might be the first that NFL fans get to experience exactly why.