Active Degradation Explains the Distribution of Nuclear Proteins during Cellular Senescence
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{"title"=>"Active degradation explains the distribution of nuclear proteins during cellular senescence", "type"=>"journal", "authors"=>[{"first_name"=>"Enrico", "last_name"=>"Giampieri", "scopus_author_id"=>"6507259256"}, {"first_name"=>"Marco", "last_name"=>"De Cecco", "scopus_author_id"=>"35739950800"}, {"first_name"=>"Daniel", "last_name"=>"Remondini", "scopus_author_id"=>"8988871200"}, {"first_name"=>"John", "last_name"=>"Sedivy", "scopus_author_id"=>"7004344606"}, {"first_name"=>"Gastone", "last_name"=>"Castellani", "scopus_author_id"=>"55925089300"}], "year"=>2015, "source"=>"PLoS ONE", "identifiers"=>{"scopus"=>"2-s2.0-84938409376", "sgr"=>"84938409376", "issn"=>"19326203", "pmid"=>"26115222", "doi"=>"10.1371/journal.pone.0118442", "pui"=>"605407796"}, "id"=>"a3d4fe85-3ddb-39de-830c-b7d842e304c7", "abstract"=>"The amount of cellular proteins is a crucial parameter that is known to vary between cells as a function of the replicative passages, and can be important during physiological aging. The process of protein degradation is known to be performed by a series of enzymatic reactions, ranging from an initial step of protein ubiquitination to their final fragmentation by the proteasome. In this paper we propose a stochastic dynamical model of nuclear proteins concentration resulting from a balance between a constant production of proteins and their degradation by a cooperative enzymatic reaction. The predictions of this model are compared with experimental data obtained by fluorescence measurements of the amount of nuclear proteins in murine tail fibroblast (MTF) undergoing cellular senescence. Our model provides a three-parameter stationary distribution that is in good agreement with the experimental data even during the transition to the senescent state, where the nuclear protein concentration changes abruptly. The estimation of three parameters (cooperativity, saturation threshold, and maximal velocity of the reaction), and their evolution during replicative passages shows that only the maximal velocity varies significantly. Based on our modeling we speculate the reduction of functionality of the protein degradation mechanism as a possible competitive inhibition of the proteasome.", "link"=>"http://www.mendeley.com/research/active-degradation-explains-distribution-nuclear-proteins-during-cellular-senescence", "reader_count"=>9, "reader_count_by_academic_status"=>{"Professor > Associate Professor"=>1, "Researcher"=>1, "Student > Ph. D. Student"=>2, "Student > Postgraduate"=>1, "Other"=>1, "Student > Master"=>1, "Lecturer > Senior Lecturer"=>1, "Professor"=>1}, "reader_count_by_user_role"=>{"Professor > Associate Professor"=>1, "Researcher"=>1, "Student > Ph. D. Student"=>2, "Student > Postgraduate"=>1, "Other"=>1, "Student > Master"=>1, "Lecturer > Senior Lecturer"=>1, "Professor"=>1}, "reader_count_by_subject_area"=>{"Biochemistry, Genetics and Molecular Biology"=>2, "Agricultural and Biological Sciences"=>3, "Medicine and Dentistry"=>1, "Physics and Astronomy"=>3}, "reader_count_by_subdiscipline"=>{"Medicine and Dentistry"=>{"Medicine and Dentistry"=>1}, "Physics and Astronomy"=>{"Physics and Astronomy"=>3}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>3}, "Biochemistry, Genetics and Molecular Biology"=>{"Biochemistry, Genetics and Molecular Biology"=>2}}, "reader_count_by_country"=>{"United States"=>1}, "group_count"=>1}

Scopus | Further Information

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/2153350"], "description"=>"<p>The upper graph is linearly scaled, the lower one is logarithmically scaled to show the distribution at high n. The black line is the best estimated distribution, while the gray area represents the uncertainty in the distribution.</p>", "links"=>[], "tags"=>["murine tail fibroblast", "mtf", "protein degradation mechanism", "protein concentration changes", "replicative passages", "model"], "article_id"=>1466526, "categories"=>["Uncategorised"], "users"=>["Enrico Giampieri", "Marco De Cecco", "Daniel Remondini", "John Sedivy", "Gastone Castellani"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0118442.g001", "stats"=>{"downloads"=>0, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_agreement_between_the_data_and_the_fitted_distribution_for_the_third_passage_/1466526", "title"=>"The agreement between the data and the fitted distribution for the third passage.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-06-26 05:01:30"}
  • {"files"=>["https://ndownloader.figshare.com/files/2153355"], "description"=>"<p>The upper graph is linearly scaled, the lower one is logarithmically scaled to show the distribution at high n. The black line is the best estimated distribution, while the gray area represents the uncertainty in the distribution.</p>", "links"=>[], "tags"=>["murine tail fibroblast", "mtf", "protein degradation mechanism", "protein concentration changes", "replicative passages", "model"], "article_id"=>1466530, "categories"=>["Uncategorised"], "users"=>["Enrico Giampieri", "Marco De Cecco", "Daniel Remondini", "John Sedivy", "Gastone Castellani"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0118442.g002", "stats"=>{"downloads"=>0, "page_views"=>1, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_agreement_between_the_data_and_the_fitted_distribution_for_the_ninth_passage_/1466530", "title"=>"The agreement between the data and the fitted distribution for the ninth passage.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-06-26 05:01:30"}
  • {"files"=>["https://ndownloader.figshare.com/files/2153356"], "description"=>"<p>The upper graph is linearly scaled, the lower one is logarithmically scaled to show the distribution at high n. The black line is the best estimated distribution, while the gray area represents the uncertainty in the distribution.</p>", "links"=>[], "tags"=>["murine tail fibroblast", "mtf", "protein degradation mechanism", "protein concentration changes", "replicative passages", "model"], "article_id"=>1466531, "categories"=>["Uncategorised"], "users"=>["Enrico Giampieri", "Marco De Cecco", "Daniel Remondini", "John Sedivy", "Gastone Castellani"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0118442.g003", "stats"=>{"downloads"=>1, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_agreement_between_the_data_and_the_fitted_distribution_for_the_tenth_passage_/1466531", "title"=>"The agreement between the data and the fitted distribution for the tenth passage.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-06-26 05:01:30"}
  • {"files"=>["https://ndownloader.figshare.com/files/2153358"], "description"=>"<p>The upper graph is linearly scaled, the lower one is logarithmically scaled to show the distribution at high n. The black line is the best estimated distribution, while the gray area represents the uncertainty in the distribution.</p>", "links"=>[], "tags"=>["murine tail fibroblast", "mtf", "protein degradation mechanism", "protein concentration changes", "replicative passages", "model"], "article_id"=>1466533, "categories"=>["Uncategorised"], "users"=>["Enrico Giampieri", "Marco De Cecco", "Daniel Remondini", "John Sedivy", "Gastone Castellani"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0118442.g004", "stats"=>{"downloads"=>0, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_agreement_between_the_data_and_the_fitted_distribution_for_the_eleventh_passage_/1466533", "title"=>"The agreement between the data and the fitted distribution for the eleventh passage.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-06-26 05:01:30"}
  • {"files"=>["https://ndownloader.figshare.com/files/2153359"], "description"=>"<p>The upper graph is linearly scaled, the lower one is logarithmically scaled to show the distribution at high n. The black line is the best estimated distribution, while the gray area represents the uncertainty in the distribution.</p>", "links"=>[], "tags"=>["murine tail fibroblast", "mtf", "protein degradation mechanism", "protein concentration changes", "replicative passages", "model"], "article_id"=>1466534, "categories"=>["Uncategorised"], "users"=>["Enrico Giampieri", "Marco De Cecco", "Daniel Remondini", "John Sedivy", "Gastone Castellani"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0118442.g005", "stats"=>{"downloads"=>0, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_agreement_between_the_data_and_the_fitted_distribution_for_the_twelfth_passage_/1466534", "title"=>"The agreement between the data and the fitted distribution for the twelfth passage.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-06-26 05:01:30"}
  • {"files"=>["https://ndownloader.figshare.com/files/2153361"], "description"=>"<p>The upper graph is linearly scaled, the lower one is logarithmically scaled to show the distribution at high n. The black line is the best estimated distribution, while the gray area represents the uncertainty in the distribution.</p>", "links"=>[], "tags"=>["murine tail fibroblast", "mtf", "protein degradation mechanism", "protein concentration changes", "replicative passages", "model"], "article_id"=>1466536, "categories"=>["Uncategorised"], "users"=>["Enrico Giampieri", "Marco De Cecco", "Daniel Remondini", "John Sedivy", "Gastone Castellani"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0118442.g006", "stats"=>{"downloads"=>1, "page_views"=>15, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_agreement_between_the_data_and_the_fitted_distribution_for_the_thirthinth_passage_/1466536", "title"=>"The agreement between the data and the fitted distribution for the thirthinth passage.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-06-26 05:01:30"}
  • {"files"=>["https://ndownloader.figshare.com/files/2153366"], "description"=>"<p>We can see that the parameters <i>α</i> and <i>θ</i> are compatible with a constant value, while the parameter <i>γ</i>, here shown as the log odd of the value, is compatible with a decrease during replicative senescence. The p value of the hypothesis of no variation during passages is 0.443 for <i>α</i>, 0.832 for <i>θ</i> and 0.048 for <i>γ</i>, evaluated with a weighted least square regression. The only parameter to varies significantly is thus <i>γ</i>, the balance between the protein creation and destruction rate.</p>", "links"=>[], "tags"=>["murine tail fibroblast", "mtf", "protein degradation mechanism", "protein concentration changes", "replicative passages", "model"], "article_id"=>1466541, "categories"=>["Uncategorised"], "users"=>["Enrico Giampieri", "Marco De Cecco", "Daniel Remondini", "John Sedivy", "Gastone Castellani"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0118442.g007", "stats"=>{"downloads"=>0, "page_views"=>4, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_behavior_of_the_parameters_over_time_/1466541", "title"=>"The behavior of the parameters over time.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-06-26 05:01:30"}
  • {"files"=>["https://ndownloader.figshare.com/files/2153367"], "description"=>"<p>All of them exhibit a transition in the observed value around the eleventh passage. The gray ares are the 50% and 95% confidence interval for the fit with a logistic function with four parameters: maximum and minimum value, steepness and transition point.</p>", "links"=>[], "tags"=>["murine tail fibroblast", "mtf", "protein degradation mechanism", "protein concentration changes", "replicative passages", "model"], "article_id"=>1466542, "categories"=>["Uncategorised"], "users"=>["Enrico Giampieri", "Marco De Cecco", "Daniel Remondini", "John Sedivy", "Gastone Castellani"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0118442.g008", "stats"=>{"downloads"=>1, "page_views"=>15, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_progression_in_time_of_three_different_centrality_measures_of_the_fitted_distribution_mode_mean_and_median_/1466542", "title"=>"The progression in time of three different centrality measures of the fitted distribution: mode, mean and median.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-06-26 05:01:30"}
  • {"files"=>["https://ndownloader.figshare.com/files/2153368"], "description"=>"<p>In the first line we have the distribution of each parameter. In the second line we have the joint probability distribution of each couple. This show that the values are correlated but without strong non–linearity.</p>", "links"=>[], "tags"=>["murine tail fibroblast", "mtf", "protein degradation mechanism", "protein concentration changes", "replicative passages", "model"], "article_id"=>1466543, "categories"=>["Uncategorised"], "users"=>["Enrico Giampieri", "Marco De Cecco", "Daniel Remondini", "John Sedivy", "Gastone Castellani"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0118442.g009", "stats"=>{"downloads"=>1, "page_views"=>7, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_distribution_of_the_parameters_945_952_and_947_for_the_third_passage_estimated_with_bayesian_analysis_/1466543", "title"=>"The distribution of the parameters (<i>α</i>, <i>θ</i> and <i>γ</i>) for the third passage estimated with bayesian analysis.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-06-26 05:01:30"}
  • {"files"=>["https://ndownloader.figshare.com/files/2153370"], "description"=>"<p>In the first line we have the distribution of each parameter. In the second line we have the joint probability distribution of each couple. This show that the values are correlated but without strong non–linearity.</p>", "links"=>[], "tags"=>["murine tail fibroblast", "mtf", "protein degradation mechanism", "protein concentration changes", "replicative passages", "model"], "article_id"=>1466545, "categories"=>["Uncategorised"], "users"=>["Enrico Giampieri", "Marco De Cecco", "Daniel Remondini", "John Sedivy", "Gastone Castellani"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0118442.g010", "stats"=>{"downloads"=>0, "page_views"=>6, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_distribution_of_the_parameters_945_952_and_947_for_the_ninth_passage_estimated_with_bayesian_analysis_/1466545", "title"=>"The distribution of the parameters (<i>α</i>, <i>θ</i> and <i>γ</i>) for the ninth passage estimated with bayesian analysis.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-06-26 05:01:30"}
  • {"files"=>["https://ndownloader.figshare.com/files/2153372"], "description"=>"<p>In the first line we have the distribution of each parameter. In the second line we have the joint probability distribution of each couple. This show that the values are correlated but without strong non–linearity.</p>", "links"=>[], "tags"=>["murine tail fibroblast", "mtf", "protein degradation mechanism", "protein concentration changes", "replicative passages", "model"], "article_id"=>1466547, "categories"=>["Uncategorised"], "users"=>["Enrico Giampieri", "Marco De Cecco", "Daniel Remondini", "John Sedivy", "Gastone Castellani"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0118442.g011", "stats"=>{"downloads"=>4, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_distribution_of_the_parameters_945_952_and_947_for_the_tenth_passage_estimated_with_bayesian_analysis_/1466547", "title"=>"The distribution of the parameters (<i>α</i>, <i>θ</i> and <i>γ</i>) for the tenth passage estimated with bayesian analysis.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-06-26 05:01:30"}
  • {"files"=>["https://ndownloader.figshare.com/files/2153373"], "description"=>"<p>In the first line we have the distribution of each parameter. In the second line we have the joint probability distribution of each couple. This show that the values are correlated but without strong non–linearity.</p>", "links"=>[], "tags"=>["murine tail fibroblast", "mtf", "protein degradation mechanism", "protein concentration changes", "replicative passages", "model"], "article_id"=>1466548, "categories"=>["Uncategorised"], "users"=>["Enrico Giampieri", "Marco De Cecco", "Daniel Remondini", "John Sedivy", "Gastone Castellani"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0118442.g012", "stats"=>{"downloads"=>0, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_distribution_of_the_parameters_945_952_and_947_for_the_eleventh_passage_estimated_with_bayesian_analysis_/1466548", "title"=>"The distribution of the parameters (<i>α</i>, <i>θ</i> and <i>γ</i>) for the eleventh passage estimated with bayesian analysis.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-06-26 05:01:30"}
  • {"files"=>["https://ndownloader.figshare.com/files/2153376"], "description"=>"<p>In the first line we have the distribution of each parameter. In the second line we have the joint probability distribution of each couple. This show that the values are correlated but without strong non–linearity.</p>", "links"=>[], "tags"=>["murine tail fibroblast", "mtf", "protein degradation mechanism", "protein concentration changes", "replicative passages", "model"], "article_id"=>1466551, "categories"=>["Uncategorised"], "users"=>["Enrico Giampieri", "Marco De Cecco", "Daniel Remondini", "John Sedivy", "Gastone Castellani"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0118442.g013", "stats"=>{"downloads"=>0, "page_views"=>6, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_distribution_of_the_parameters_945_952_and_947_for_the_twelfth_passage_estimated_with_bayesian_analysis_/1466551", "title"=>"The distribution of the parameters (<i>α</i>, <i>θ</i> and <i>γ</i>) for the twelfth passage estimated with bayesian analysis.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-06-26 05:01:30"}
  • {"files"=>["https://ndownloader.figshare.com/files/2153379"], "description"=>"<p>In the first line we have the distribution of each parameter. In the second line we have the joint probability distribution of each couple. This show that the values are correlated but without strong non–linearity.</p>", "links"=>[], "tags"=>["murine tail fibroblast", "mtf", "protein degradation mechanism", "protein concentration changes", "replicative passages", "model"], "article_id"=>1466554, "categories"=>["Uncategorised"], "users"=>["Enrico Giampieri", "Marco De Cecco", "Daniel Remondini", "John Sedivy", "Gastone Castellani"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0118442.g014", "stats"=>{"downloads"=>1, "page_views"=>11, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_distribution_of_the_parameters_945_952_and_947_for_the_thirthinth_passage_estimated_with_bayesian_analysis_/1466554", "title"=>"The distribution of the parameters (<i>α</i>, <i>θ</i> and <i>γ</i>) for the thirthinth passage estimated with bayesian analysis.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2015-06-26 05:01:30"}
  • {"files"=>["https://ndownloader.figshare.com/files/2153380"], "description"=>"<p>Each row represent a different experiment, that was evaluated individually. The row labeled as <i>combined</i><i>experiments</i> is the result of the fit using all the values from all experiments for each passage (where available) to increase the size of the sample. The p value represent the hypothesys that the data comes from the distribution, so low p values indicate poor agreement. For most cases the negative binomial fit poorly the data (low p values) while the generalized version perform correctly in all cases (all p values are above 0.05).</p>", "links"=>[], "tags"=>["murine tail fibroblast", "mtf", "protein degradation mechanism", "protein concentration changes", "replicative passages", "model"], "article_id"=>1466555, "categories"=>["Uncategorised"], "users"=>["Enrico Giampieri", "Marco De Cecco", "Daniel Remondini", "John Sedivy", "Gastone Castellani"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0118442.t001", "stats"=>{"downloads"=>1, "page_views"=>3, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_p_values_for_the_fit_between_the_described_distributions_and_the_data_/1466555", "title"=>"<i>p values</i> for the fit between the described distributions and the data.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2015-06-26 05:01:30"}
  • {"files"=>["https://ndownloader.figshare.com/files/2153381"], "description"=>"<p>A negative value imply a preference toward the generalized negative binomial. The generalized negative binomial is preferred in all cases for both measurements, aside for the BIC value of the tenth passage where the difference in close to 0 (so they are equivalent). Bigger dataset (passages 3 and 13) evidence a strong preference toward the generalized negative binomial. These results are robust under correction for small sample size (that gives a correction of order 10<sup>−1</sup>). As BIC penalizes strongly the higher number of parameters of the generalized negative binomial we obtain values lower than those of the AIC, but with the same general trend.</p>", "links"=>[], "tags"=>["murine tail fibroblast", "mtf", "protein degradation mechanism", "protein concentration changes", "replicative passages", "model"], "article_id"=>1466556, "categories"=>["Uncategorised"], "users"=>["Enrico Giampieri", "Marco De Cecco", "Daniel Remondini", "John Sedivy", "Gastone Castellani"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0118442.t002", "stats"=>{"downloads"=>6, "page_views"=>13, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_The_AIC_and_BIC_differences_between_the_two_models_/1466556", "title"=>"The AIC and BIC differences between the two models.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2015-06-26 05:01:30"}
  • {"files"=>["https://ndownloader.figshare.com/files/2153382"], "description"=>"<div><p>The amount of cellular proteins is a crucial parameter that is known to vary between cells as a function of the replicative passages, and can be important during physiological aging. The process of protein degradation is known to be performed by a series of enzymatic reactions, ranging from an initial step of protein ubiquitination to their final fragmentation by the proteasome. In this paper we propose a stochastic dynamical model of nuclear proteins concentration resulting from a balance between a constant production of proteins and their degradation by a cooperative enzymatic reaction. The predictions of this model are compared with experimental data obtained by fluorescence measurements of the amount of nuclear proteins in murine tail fibroblast (MTF) undergoing cellular senescence. Our model provides a three-parameter stationary distribution that is in good agreement with the experimental data even during the transition to the senescent state, where the nuclear protein concentration changes abruptly. The estimation of three parameters (cooperativity, saturation threshold, and maximal velocity of the reaction), and their evolution during replicative passages shows that only the maximal velocity varies significantly. Based on our modeling we speculate the reduction of functionality of the protein degradation mechanism as a possible competitive inhibition of the proteasome.</p></div>", "links"=>[], "tags"=>["murine tail fibroblast", "mtf", "protein degradation mechanism", "protein concentration changes", "replicative passages", "model"], "article_id"=>1466557, "categories"=>["Uncategorised"], "users"=>["Enrico Giampieri", "Marco De Cecco", "Daniel Remondini", "John Sedivy", "Gastone Castellani"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0118442", "stats"=>{"downloads"=>2, "page_views"=>9, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Active_Degradation_Explains_the_Distribution_of_Nuclear_Proteins_during_Cellular_Senescence_/1466557", "title"=>"Active Degradation Explains the Distribution of Nuclear Proteins during Cellular Senescence", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2015-06-26 05:01:30"}

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{"start_date"=>"2015-01-01T00:00:00Z", "end_date"=>"2015-12-31T00:00:00Z", "subject_areas"=>[]}
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