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Early anatomical studies suggested that the VL is organized in a \u201cfan\u2010out fan\u2010in\u201d connectivity matrix comprising only three morphologically identified neuron types; input axons from the median superior frontal lobe (MSFL) innervating\n                    <jats:italic>en passant<\/jats:italic>\n                    millions of small amacrine interneurons (AMs), which converge sharply onto large VL output neurons (LNs). Recent physiological studies confirmed the feedforward excitatory connectivity; a glutamatergic synapse at the first MSFL\u2010to\u2010AM synaptic layer and a cholinergic AM\u2010to\u2010LNs synapse. MSFL\u2010to\u2010AMs synapses show a robust hippocampal\u2010like activity\u2010dependent long\u2010term potentiation (LTP) of transmitter release. 5\u2010HT, octopamine, dopamine and nitric oxide modulate short\u2010 and long\u2010term VL synaptic plasticity. Here, we present a comprehensive histolabeling study to better characterize the neural elements in the VL. We generally confirmed glutamatergic MSFLs and cholinergic AMs. Intense labeling for NOS activity in the AMs neurites were in\u2010line with the NO\u2010dependent presynaptic LTP mechanism at the MSFL\u2010to\u2010AM synapse. New discoveries here reveal more heterogeneity of the VL neurons than previously thought. GABAergic AMs suggest a subpopulation of inhibitory interneurons in the first input layer. Clear \u03b3\u2010amino butyric acid labeling in the cell bodies of LNs supported an inhibitory VL output, yet the LNs co\u2010expressed FMRFamide\u2010like neuropeptides, suggesting an additional neuromodulatory role of the VL output. Furthermore, a group of LNs was glutamatergic. A new cluster of cells organized as a \u201cdeep nucleus\u201d showed rich catecholaminergic labeling and may play a role in intrinsic neuromodulation. In\u2010situ hybridization and immunolabeling allowed characterization and localization of a rich array of neuropeptides and neuromodulators, likely involved in reward\/punishment signals. This analysis of the fast transmission system, together with the newly found cellular elements, help integrate behavioral, physiological, pharmacological and connectome findings into a more comprehensive understanding of an efficient learning and memory network.\n                  <\/jats:p>","DOI":"10.1002\/jmor.21459","type":"journal-article","created":{"date-parts":[[2022,2,2]],"date-time":"2022-02-02T08:59:47Z","timestamp":1643792387000},"page":"557-584","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":13,"title":["Neurotransmission and neuromodulation systems in the learning and memory network of\n                    <i>Octopus vulgaris<\/i>"],"prefix":"10.1002","volume":"283","author":[{"given":"Naama","family":"Stern\u2010Mentch","sequence":"first","affiliation":[{"name":"Department of Neurobiology, Silberman Institute of Life Sciences Hebrew University  Jerusalem Israel"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gabrielle Winters","family":"Bostwick","sequence":"additional","affiliation":[{"name":"Department of Neuroscience and McKnight Brain Institute, and Whitney Laboratory for Marine Bioscience University of Florida  Gainesville Florida USA"},{"name":"Ocean Genome Atlas Project  San Francisco USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michael","family":"Belenky","sequence":"additional","affiliation":[{"name":"Department of Neurobiology, Silberman Institute of Life Sciences Hebrew University  Jerusalem Israel"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1333-3176","authenticated-orcid":false,"given":"Leonid","family":"Moroz","sequence":"additional","affiliation":[{"name":"Department of Neuroscience and McKnight Brain Institute, and Whitney Laboratory for Marine Bioscience University of Florida  Gainesville Florida USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9638-7320","authenticated-orcid":false,"given":"Binyamin","family":"Hochner","sequence":"additional","affiliation":[{"name":"Department of Neurobiology, Silberman Institute of Life Sciences Hebrew University  Jerusalem Israel"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2022,2,8]]},"reference":[{"key":"e_1_2_10_2_1","doi-asserted-by":"publisher","DOI":"10.4056\/sigs.3136559"},{"key":"e_1_2_10_3_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev-animal-021419-083609"},{"key":"e_1_2_10_4_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature14668"},{"key":"e_1_2_10_5_1","doi-asserted-by":"publisher","DOI":"10.1007\/s10339-007-0192-9"},{"key":"e_1_2_10_6_1","doi-asserted-by":"crossref","unstructured":"Amodio P. &Fiorito G.(2013).Chapter 23 \u2010 Observational and Other Types of Learning in Octopus. In R. Menzel & P. R. Benjamin (Eds.) Handbook of Behavioral Neuroscience Vol. 22 pp.293\u2013302. Elsevier.https:\/\/doi.org\/10.1016\/B978-0-12-415823-8.00023-X","DOI":"10.1016\/B978-0-12-415823-8.00023-X"},{"key":"e_1_2_10_7_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.2357-07.2007"},{"key":"e_1_2_10_8_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0092-8674(00)81797-3"},{"key":"e_1_2_10_9_1","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.1990.sp018249"},{"key":"e_1_2_10_10_1","unstructured":"Bidel F. Meirovitch Y. Schalek L. Xiaotang L. Yang A. Berger D. Peleg A. Shaked A. Flash T. Lichtman J. &Hochner B. (2021).Insight into the connectome of the learning and memory network of the Octopus. Program No. P856.06. 2021 Neuroscience Meeting Planner. Society for Neuroscience"},{"key":"e_1_2_10_11_1","first-page":"157","article-title":"A review of simultaneous visual discrimination as a method of training octopuses","volume":"71","author":"Boal J. 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