{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,16]],"date-time":"2026-04-16T14:51:37Z","timestamp":1776351097840,"version":"3.51.2"},"reference-count":41,"publisher":"Springer Science and Business Media LLC","issue":"3","license":[{"start":{"date-parts":[[2012,11,22]],"date-time":"2012-11-22T00:00:00Z","timestamp":1353542400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/2.0"},{"start":{"date-parts":[[2012,11,22]],"date-time":"2012-11-22T00:00:00Z","timestamp":1353542400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/2.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["J Comput Neurosci"],"published-print":{"date-parts":[[2013,6]]},"abstract":"<jats:title>Abstract<\/jats:title>\n          <jats:p>Optogenetics offers an unprecedented ability to spatially target neuronal stimulations. This study investigated via simulation, for the first time, how the spatial pattern of excitation affects the response of channelrhodopsin-2 (ChR2) expressing neurons. First we described a methodology for modeling ChR2 in the NEURON simulation platform. Then, we compared four most commonly considered illumination strategies (somatic, dendritic, axonal and whole cell) in a paradigmatic model of a cortical layer V pyramidal cell. We show that the spatial pattern of illumination has an important impact on the efficiency of stimulation and the kinetics of the spiking output. Whole cell illumination synchronizes the depolarization of the dendritic tree and the soma and evokes spiking characteristics with a distinct pattern including an increased bursting rate and enhanced back propagation of action potentials (bAPs). This type of illumination is the most efficient as a given irradiance threshold was achievable with only 6 % of ChR2 density needed in the case of somatic illumination. Targeting only the axon initial segment requires a high ChR2 density to achieve a given threshold irradiance and a prolonged illumination does not yield sustained spiking. We also show that patterned illumination can be used to modulate the bAPs and hence spatially modulate the direction and amplitude of spike time dependent plasticity protocols. We further found the irradiance threshold to increase in proportion to the demyelination level of an axon, suggesting that measurements of the irradiance threshold (for example relative to the soma) could be used to remotely probe a loss of neural myelin sheath, which is a hallmark of several neurodegenerative diseases.<\/jats:p>","DOI":"10.1007\/s10827-012-0431-7","type":"journal-article","created":{"date-parts":[[2012,11,22]],"date-time":"2012-11-22T17:47:37Z","timestamp":1353606457000},"page":"477-488","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":52,"title":["The spatial pattern of light determines the kinetics and modulates backpropagation of optogenetic action potentials"],"prefix":"10.1007","volume":"34","author":[{"given":"Nir","family":"Grossman","sequence":"first","affiliation":[]},{"given":"Vasiliki","family":"Simiaki","sequence":"additional","affiliation":[]},{"given":"Claire","family":"Martinet","sequence":"additional","affiliation":[]},{"given":"Christofer","family":"Toumazou","sequence":"additional","affiliation":[]},{"given":"Simon R.","family":"Schultz","sequence":"additional","affiliation":[]},{"given":"Konstantin","family":"Nikolic","sequence":"additional","affiliation":[]}],"member":"297","published-online":{"date-parts":[[2012,11,22]]},"reference":[{"issue":"26","key":"431_CR1","doi-asserted-by":"publisher","first-page":"11981","DOI":"10.1073\/pnas.1006620107","volume":"107","author":"BK Andrasfalvy","year":"2010","unstructured":"Andrasfalvy, B.K., Zemelman, B.V., Tang, J., Vaziri, A. (2010). Two-photon single-cell optogenetic control of neuronal activity by sculpted light. Proceedings of the National Academy of Sciences of the United States of America, 107(26), 11981\u201311986.","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"key":"431_CR2","first-page":"13","volume-title":"Optical Control of Neural Excitability","author":"E Bamberg","year":"2008","unstructured":"Bamberg, E., Bamann, C., Feldbauer, K., Kleinlogel, S., Spitz, J., Zimmermann, D., Wood, P., Nagel, G. (2008). Channelrhodopsins: Molecular properties and applications. In H. Keshishian (Ed.), Optical Control of Neural Excitability, (pp. 13\u201320). Washington, DC: Society for Neuroscience."},{"issue":"9","key":"431_CR3","doi-asserted-by":"publisher","first-page":"1263","DOI":"10.1038\/nn1525","volume":"8","author":"ES Boyden","year":"2005","unstructured":"Boyden, E.S., Zhang, F., Bamberg, E., Nagel, G., Deisseroth, K. (2005). Millisecond-timescale, genetically targeted optical control of neural activity. Nature Neuroscience, 8(9), 1263\u20131268.","journal-title":"Nature Neuroscience"},{"key":"431_CR4","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.conb.2011.06.001","volume":"21","author":"V Busskamp","year":"2011","unstructured":"Busskamp, V., & Roska, B. (2011). Optogenetic approaches to restoring visual function in retinitis pigmentosa. Current Opinion in Neurobiology, 21, 1\u20135.","journal-title":"Current Opinion in Neurobiology"},{"key":"431_CR5","doi-asserted-by":"publisher","DOI":"10.1017\/CBO9780511541612","volume-title":"The neuron book","author":"N Carnevale","year":"2006","unstructured":"Carnevale, N., & Hines, M. (2006). The neuron book. Cambridge, UK: Cambridge University Press."},{"issue":"7277","key":"431_CR6","doi-asserted-by":"publisher","first-page":"98","DOI":"10.1038\/nature08652","volume":"463","author":"BY Chow","year":"2010","unstructured":"Chow, B.Y., Han, X., Dobry, A.S., Qian, X., Chuong, A.S., Li, M., Henninger, M.A., Belfort, G.M., Lin, Y., Monahan, P.E., Boyden, E.S. (2010). High-performance genetically targetable optical neural silencing by light-driven proton pumps. Nature, 463(7277), 98\u2013102.","journal-title":"Nature"},{"issue":"3","key":"431_CR7","doi-asserted-by":"publisher","first-page":"035007","DOI":"10.1088\/1741-2560\/6\/3\/035007","volume":"6","author":"P Degenaar","year":"2009","unstructured":"Degenaar, P., Grossman, N., Memon, M., Burrone, J., Dawson, M., Drakakis, E., Neil, M., Nikolic, K. (2009). Optobionic vision\u2014a new genetically enhanced light on retinal prosthesis. Journal of Neural Engineering, 6(3), 035007.","journal-title":"Journal of Neural Engineering"},{"issue":"30","key":"431_CR8","doi-asserted-by":"publisher","first-page":"12317","DOI":"10.1073\/pnas.0905852106","volume":"106","author":"K Feldbauer","year":"2009","unstructured":"Feldbauer, K., Zimmermann, D., Pintschovius, V., Spitz, J., Bamann, C., Bamberg, E. (2009). Channelrhodopsin-2 is a leaky proton pump. Proceedings of the National Academy of Sciences of the United States of America, 106(30), 12317\u201312322.","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"issue":"6","key":"431_CR9","doi-asserted-by":"publisher","first-page":"1742","DOI":"10.1109\/TBME.2011.2114883","volume":"58","author":"N Grossman","year":"2011","unstructured":"Grossman, N., Nikolic, K., Toumazou, C., Degenaar, P. (2011). Modeling study of the light stimulation of a neuron cell with channelrhodopsin-2 mutants. IEEE Transactions on Biomedical Engineering, 58(6), 1742\u20131751.","journal-title":"IEEE Transactions on Biomedical Engineering"},{"issue":"1","key":"431_CR10","doi-asserted-by":"publisher","first-page":"016004","DOI":"10.1088\/1741-2560\/7\/1\/016004","volume":"7","author":"N Grossman","year":"2010","unstructured":"Grossman, N., Poher, P., Grubb, M.S., Kennedy, G.T., Nikolic, K., McGovern, B., Berlinguer Palmini, R., Gong, Z., Drakakis, E.M., Neil, M.A.A., Dawson, M.D., Burrone, J., Degenaar, P. (2010). Multi-site optical excitation using chr2 and micro-led array. Journal of Neural Engineering, 7(1), 016004.","journal-title":"Journal of Neural Engineering"},{"issue":"3","key":"431_CR11","doi-asserted-by":"publisher","first-page":"387","DOI":"10.1038\/nn.2495","volume":"13","author":"LA Gunaydin","year":"2010","unstructured":"Gunaydin, L.A., Yizhar, O., Berndt, A., Sohal, V.S., Deisseroth, K., Hegemann, P. (2010). Ultrafast optogenetic control. Nature Neuroscience, 13(3), 387\u201392.","journal-title":"Nature Neuroscience"},{"issue":"2","key":"431_CR12","doi-asserted-by":"publisher","first-page":"191","DOI":"10.1016\/j.neuron.2009.03.011","volume":"62","author":"X Han","year":"2009","unstructured":"Han, X., Qian, X., Bernstein, J.G., Zhou, H.H., Franzesi, G.T., Stern, P., Bronson, R.T., Graybiel, A.M., Desimone, R., Boyden, E.S. (2009). Millisecond-timescale optical control of neural dynamics in the nonhuman primate brain. Neuron, 62(2), 191\u2013198.","journal-title":"Neuron"},{"issue":"7","key":"431_CR13","doi-asserted-by":"publisher","first-page":"e1002107","DOI":"10.1371\/journal.pcbi.1002107","volume":"7","author":"E Hay","year":"2011","unstructured":"Hay, E., Hill, S., Schrmann, F., Markram, H., Segev, I. (2011). Models of neocortical layer 5b pyramidal cells capturing awide range of dendritic and perisomatic active properties. PLoS Computational Biology, 7(7), e1002107.","journal-title":"PLoS Computational Biology"},{"issue":"6","key":"431_CR14","doi-asserted-by":"publisher","first-page":"3911","DOI":"10.1529\/biophysj.105.069716","volume":"89","author":"P Hegemann","year":"2005","unstructured":"Hegemann, P., Ehlenbeck, S., Gradmann, D. (2005). Multiple photocycles of channelrhodopsin. Biophysical Journal, 89(6), 3911\u20133918.","journal-title":"Biophysical Journal"},{"key":"431_CR15","doi-asserted-by":"publisher","first-page":"1179","DOI":"10.1162\/neco.1997.9.6.1179","volume":"9","author":"M Hines","year":"1997","unstructured":"Hines, M., & Carnevale, N. (1997). The neuron simulation environment. Neural Computation, 9, 1179\u20131209.","journal-title":"Neural Computation"},{"issue":"5","key":"431_CR16","doi-asserted-by":"publisher","first-page":"995","DOI":"10.1162\/089976600300015475","volume":"12","author":"M Hines","year":"2000","unstructured":"Hines, M., & Carnevale, N. (2000). Expanding neuron\u2019s repertoire of mechanisms with nmodl. Neural Computation, 12(5), 995\u20131007.","journal-title":"Neural Computation"},{"issue":"8","key":"431_CR17","doi-asserted-by":"publisher","first-page":"996","DOI":"10.1038\/nn.2359","volume":"12","author":"W Hu","year":"2009","unstructured":"Hu, W., Tian, C., Li, T., Yang, M., Hou, H., Shu, Y. (2009). Distinct contributions of nav1.6 and nav1.2 in action potential initiation and backpropagation. Nature Neuroscience, 12(8), 996\u20131002.","journal-title":"Nature Neuroscience"},{"issue":"2","key":"431_CR18","doi-asserted-by":"publisher","first-page":"178","DOI":"10.1038\/nn2040","volume":"11","author":"MHP Kole","year":"2008","unstructured":"Kole, M.H.P., Ilschner, S.U., Kampa, B.M., Williams, S.R., Ruben, P.C., Stuart, G.J. (2008). Action potential generation requires a high sodium channel density in the axon initial segment. Nature Neuroscience, 11(2), 178\u2013186.","journal-title":"Nature Neuroscience"},{"issue":"41","key":"431_CR19","doi-asserted-by":"publisher","first-page":"10420","DOI":"10.1523\/JNEUROSCI.2650-06.2006","volume":"26","author":"JJ Letzkus","year":"2006","unstructured":"Letzkus, J.J., Kampa, B.M., Stuart, G.J. (2006). Learning rules for spike timing-dependent plasticity depend on dendritic synapse location. The Journal of Neuroscience, 26(41), 10420\u201310429.","journal-title":"The Journal of Neuroscience"},{"issue":"5","key":"431_CR20","doi-asserted-by":"publisher","first-page":"1803","DOI":"10.1016\/j.bpj.2008.11.034","volume":"96","author":"JY Lin","year":"2009","unstructured":"Lin, J.Y., Lin, M.Z., Steinbach, P., Tsien, R.Y. (2009). Characterization of engineered channelrhodopsin variants with improved properties and kinetics. Biophysical Journal, 96(5), 1803\u20131814.","journal-title":"Biophysical Journal"},{"issue":"9","key":"431_CR21","doi-asserted-by":"publisher","first-page":"821","DOI":"10.1038\/nmeth.1241","volume":"5","author":"C Lutz","year":"2008","unstructured":"Lutz, C., Otis, T.S., DeSars, V., Charpak, S., DiGregorio, D.A., Emiliani, V. (2008). Holographic photolysis of caged neurotransmitters. Nature Methods, 5(9), 821\u2013827.","journal-title":"Nature Methods"},{"key":"431_CR22","doi-asserted-by":"publisher","first-page":"363","DOI":"10.1038\/382363a0","volume":"382","author":"ZF Mainen","year":"1996","unstructured":"Mainen, Z.F., & Sejnowski, T.J. (1996). Influence of dendritic structure on firing pattern in model neocortical neurons. Nature, 382, 363\u2013366.","journal-title":"Nature"},{"issue":"24","key":"431_CR23","doi-asserted-by":"publisher","first-page":"13940","DOI":"10.1073\/pnas.1936192100","volume":"100","author":"G Nagel","year":"2003","unstructured":"Nagel, G., Szellas, T., Huhn, W., Kateriya, S., Adeishvili, N., Berthold, P., Ollig, D., Hegemann, P., Bamberg, E. (2003). Channelrhodopsin-2, a directly light-gated cation-selective membrane channel. Proceedings of the National Academy of Sciences of the United States of America, 100(24), 13940\u201313945.","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"key":"431_CR24","first-page":"1626","volume":"1","author":"K Nikolic","year":"2006","unstructured":"Nikolic, K., Degenaar, P., Toumazou, C. (2006). Modeling and engineering aspects of channelrhodopsin2 system for neural photostimulation. Conference Proceedings IEEE EMBS, 1, 1626\u20131629.","journal-title":"Conference Proceedings IEEE EMBS"},{"key":"431_CR25","doi-asserted-by":"publisher","first-page":"400","DOI":"10.1111\/j.1751-1097.2008.00460.x","volume":"85","author":"K Nikolic","year":"2009","unstructured":"Nikolic, K., Grossman, N., Grubb, M.S., Burrone, J., Toumazou, C., Degenaar, P. (2009). Photocycles of channelrhodopsin-2. Photochemistry and Photobiology, 85, 400\u2013411.","journal-title":"Photochemistry and Photobiology"},{"issue":"10","key":"431_CR26","doi-asserted-by":"publisher","first-page":"848","DOI":"10.1038\/nmeth.1505","volume":"7","author":"E Papagiakoumou","year":"2010","unstructured":"Papagiakoumou, E., Anselmi, F., Begue, A., de Sars, V., Gluckstad, J., Isacoff, E.Y., Emiliani, V. (2010). Scanless two-photon excitation of channelrhodopsin-2. Nature Methods, 7(10), 848\u2013854.","journal-title":"Nature Methods"},{"issue":"35","key":"431_CR27","doi-asserted-by":"publisher","first-page":"15025","DOI":"10.1073\/pnas.0907084106","volume":"106","author":"JP Rickgauer","year":"2009","unstructured":"Rickgauer, J.P., & Tank, D.W. (2009). Two-photon excitation of channelrhodopsin-2 at saturation. Proceedings of the National Academy of Sciences, 106(35), 15025\u201315030.","journal-title":"Proceedings of the National Academy of Sciences"},{"issue":"6","key":"431_CR28","doi-asserted-by":"publisher","first-page":"3143","DOI":"10.1152\/jn.00046.2003","volume":"89","author":"AT Schaefer","year":"2003","unstructured":"Schaefer, A.T., Larkum, M.E., Sakmann, B., Roth, A. (2003). Coincidence detection in pyramidal neurons is tuned by their dendritic branching pattern. Journal of Neurophysiology, 89(6), 3143\u20133154.","journal-title":"Journal of Neurophysiology"},{"issue":"11","key":"431_CR29","doi-asserted-by":"publisher","first-page":"837","DOI":"10.1038\/nmeth793","volume":"2","author":"S Shoham","year":"2005","unstructured":"Shoham, S., O\u2019Connor, D.H., Sarkisov, D.V., Wang, S.S.H. (2005). Rapid neurotransmitter uncaging in spatially defined patterns. Nature Methods, 2(11), 837\u2013843.","journal-title":"Nature Methods"},{"issue":"7094","key":"431_CR30","doi-asserted-by":"publisher","first-page":"761","DOI":"10.1038\/nature04720","volume":"441","author":"Y Shu","year":"2006","unstructured":"Shu, Y., Hasenstaub, A., Duque, A., Yu, Y., McCormick, D.A. (2006). Modulation of intracortical synaptic potentials by presynaptic somatic membrane potential. Nature, 441(7094), 761\u2013765.","journal-title":"Nature"},{"key":"431_CR31","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1038\/367069a0","volume":"367","author":"GJ Stuart","year":"1994","unstructured":"Stuart, G.J., & Sakmann, B. (1994). Active propagation of somatic action potentials into neocortical pyramidal cell dendrites. Nature, 367, 69\u201372.","journal-title":"Nature"},{"issue":"3","key":"431_CR32","doi-asserted-by":"publisher","first-page":"617","DOI":"10.1111\/j.1469-7793.1997.617ba.x","volume":"505","author":"G Stuart","year":"1997","unstructured":"Stuart, G., Schiller, J., Sakmann, B. (1997). Action potential initiation and propagation in rat neocortical pyramidal neurons. The Journal of Physiology, 505(3), 617\u2013632.","journal-title":"The Journal of Physiology"},{"issue":"4","key":"431_CR33","doi-asserted-by":"publisher","first-page":"1767","DOI":"10.1016\/S0006-3495(98)77887-7","volume":"74","author":"O Tour","year":"1998","unstructured":"Tour, O., Parnas, H., Parnas, I. (1998). Depolarization increases the single-channel conductance and the open probability of crayfish glutamate channels. Biophysical Journal, 74(4), 1767\u20131778.","journal-title":"Biophysical Journal"},{"key":"431_CR34","doi-asserted-by":"publisher","first-page":"8143","DOI":"10.1073\/pnas.0700384104","volume":"104","author":"H Wang","year":"2007","unstructured":"Wang, H., Peca, J., Matsuzaki, M., Matsuzaki, K., Noguchi, J., Qiu, L., Wang, D., Zhang, F., Boyden, E., Deisseroth, K., Kasai, H., Hall, W.C., Feng, G., Augustine, G.J. (2007). High-speed mapping of synaptic connectivity using photostimulation in channelrhodopsin-2 transgenic mice. Proceedings of the National Academy of Sciences of the United States of America, 104, 8143\u20138148.","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"issue":"4","key":"431_CR35","doi-asserted-by":"publisher","first-page":"046002","DOI":"10.1088\/1741-2560\/8\/4\/046002","volume":"8","author":"S Yang","year":"2011","unstructured":"Yang, S., Papagiakoumou, E., Guillon, M., de Sars, V., Tang, C.-M., Emiliani, V. (2011). Three-dimensional holographic photostimulation of the dendritic arbor. Journal of Neural Engineering, 8(4), 046002.","journal-title":"Journal of Neural Engineering"},{"issue":"1","key":"431_CR36","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1016\/j.neuron.2011.06.004","volume":"71","author":"O Yizhar","year":"2011","unstructured":"Yizhar, O., Fenno, L., Davidson, T., Mogri, M., Deisseroth, K. (2011). Optogenetics in neural systems. Neuron, 71(1), 9\u201334.","journal-title":"Neuron"},{"issue":"29","key":"431_CR37","doi-asserted-by":"publisher","first-page":"7260","DOI":"10.1523\/JNEUROSCI.1613-08.2008","volume":"28","author":"Y Yu","year":"2008","unstructured":"Yu, Y., Shu, Y., McCormick, D.A. (2008). Cortical action potential backpropagation explains spike threshold variability and rapid-onset kinetics. Journal of Neuroscience, 28(29), 7260\u20137272.","journal-title":"Journal of Neuroscience"},{"issue":"8","key":"431_CR38","doi-asserted-by":"publisher","first-page":"577","DOI":"10.1038\/nrn2192","volume":"8","author":"F Zhang","year":"2007","unstructured":"Zhang, F., Aravanis, A.M., Adamantidis, A., de Lecea, L., Deisseroth, K. (2007a). Circuit-breakers: optical technologies for probing neural signals and systems. Nature Reviews Neuroscience, 8(8), 577\u201381.","journal-title":"Nature Reviews Neuroscience"},{"issue":"10","key":"431_CR39","doi-asserted-by":"publisher","first-page":"785","DOI":"10.1038\/nmeth936","volume":"3","author":"F Zhang","year":"2006","unstructured":"Zhang, F., Wang, L.P., Boyden, E.S., Deisseroth, K. (2006). Channelrhodopsin-2 and optical control of excitable cells. Nature Methods, 3(10), 785\u201392.","journal-title":"Nature Methods"},{"issue":"7136","key":"431_CR40","doi-asserted-by":"publisher","first-page":"633","DOI":"10.1038\/nature05744","volume":"446","author":"F Zhang","year":"2007","unstructured":"Zhang, F., Wang, L.P., Brauner, M., Liewald, J.F., Kay, K., Watzke, N., Wood, P.G., Bamberg, E., Nagel, G., Gottschalk, A., Deisseroth, K. (2007b). Multimodal fast optical interrogation of neural circuitry. Nature, 446(7136), 633\u2013639.","journal-title":"Nature"},{"issue":"2","key":"431_CR41","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1038\/nmeth988","volume":"4","author":"YP Zhang","year":"2007","unstructured":"Zhang, Y.P., & Oertner, T.G. (2007). Optical induction of synaptic plasticity using a light-sensitive channel. Nature Methods, 4(2), 139\u2013141.","journal-title":"Nature Methods"}],"container-title":["Journal of Computational Neuroscience"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s10827-012-0431-7.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s10827-012-0431-7\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"http:\/\/link.springer.com\/content\/pdf\/10.1007\/s10827-012-0431-7","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s10827-012-0431-7.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,9,1]],"date-time":"2021-09-01T20:21:09Z","timestamp":1630527669000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s10827-012-0431-7"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2012,11,22]]},"references-count":41,"journal-issue":{"issue":"3","published-print":{"date-parts":[[2013,6]]}},"alternative-id":["431"],"URL":"https:\/\/doi.org\/10.1007\/s10827-012-0431-7","relation":{},"ISSN":["0929-5313","1573-6873"],"issn-type":[{"value":"0929-5313","type":"print"},{"value":"1573-6873","type":"electronic"}],"subject":[],"published":{"date-parts":[[2012,11,22]]},"assertion":[{"value":"10 January 2012","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"21 August 2012","order":2,"name":"revised","label":"Revised","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"11 October 2012","order":3,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"22 November 2012","order":4,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}}]}}