Locating the engram: Should we look for plastic synapses or information-storing molecules?

Abdou, 2018, Synapse-specific representation of the identity of overlapping memory engrams, Science, 360, 1227, 10.1126/science.aat3810 Abraham, 2019, Is plasticity of synapses the mechanism of long-term memory storage?, NPJ Science of Learning, 4, 9, 10.1038/s41539-019-0048-y Amit, 1995, The Hebbian paradigm reintegrated: Local reverberations as internal representations, Behavioral and Brain Sciences, 18, 617, 10.1017/S0140525X00040164 Anwyl, 2009, Metabotropic glutamate receptor-dependent long-term potentiation, Neuropharmacology, 56, 735, 10.1016/j.neuropharm.2009.01.002 Aronov, 2017, Mapping of a non-spatial dimension by the hippocampal-entorhinal circuit, Nature, 543, 719, 10.1038/nature21692 Arshavsky, 2016, Neurons versus networks: The interplay between individual neurons and neural networks in cognitive functions, The Neuroscientist, 23, 341, 10.1177/1073858416670124 Attardo, 2015, Impermanence of dendritic spines in live adult CA1 hippocampus, Nature, 523, 592, 10.1038/nature14467 Balsam, 2006, Pavlovian contingencies and temporal information, Journal of Experimental Psychology: Animal Behavior Processes, 32, 284 Barot, 2009, Functional imaging of stimulus convergence in amygdalar neurons during Pavlovian fear conditioning, PLoS One, 4, e6156, 10.1371/journal.pone.0006156 Bédécarrats, 2018, RNA from trained Aplysia can induce an epigenetic engram for long-term sensitization in untrained Aplysia, eNeuro, 5, 10.1523/ENEURO.0038-18.2018 Bevins, 1997, Converging evidence for one-trial context fear conditioning with an immediate shock: Importance of shock potency, Journal of Experimental Psychology: Animal Behavior Processes, 23, 312 Brophy, 2014, Principles of genetic circuit design, Nature Methods, 101, 508, 10.1038/nmeth.2926 Brunec, 2018, Multiple scales of representation along the hippocampal anteroposterior axis in humans, Current Biology, 28, 2129, 10.1016/j.cub.2018.05.016 Buehlmann, 2014, Desert ants locate food by combining high sensitivity to food odors with extensive crosswind runs, Current Biology, 24, 960, 10.1016/j.cub.2014.02.056 Burak, 2009, Accurate path integration in continuous attractor network models of grid cells, PLoS Computational Biology, 5, 1, 10.1371/journal.pcbi.1000291 Butler, 2018, Neurons specifically activated by fear learning in lateral amygdala display increased synaptic strength, eNeuro, 5, 10.1523/ENEURO.0114-18.2018 Buzsáki, 2010, Neural syntax: Cell assemblies, synapsembles, and readers, Neuron, 68, 362, 10.1016/j.neuron.2010.09.023 Buzsáki, 2019 Carrillo-Reid, 2016, Imprinting and recalling cortical ensembles, Science, 353, 691, 10.1126/science.aaf7560 Chang, 2017, The representation of colored objects in macaque color patches, Nature Communications, 8, 2064, 10.1038/s41467-017-01912-7 Chang, 2017, The code for facial identity in the primate brain, Cell, 169, 1013, 10.1016/j.cell.2017.05.011 Chase, 2007, First-spike latency information in single neurons increases when referenced to population onset, Proceedings of the National Academy of Sciences of the United States of America, 104, 5175, 10.1073/pnas.0610368104 Chen, 2014, Reinstatement of long-term memory following erasure of its behavioral and synaptic expression in Aplysia, Elife, 3, e03896, 10.7554/eLife.03896 Choi, 2018, Interregional synaptic maps among engram cells underlie memory formation, Science, 360, 430, 10.1126/science.aas9204 Cobar, 2017, Place cells and long-term potentiation in the hippocampus, Neurobiology of Learning & Memory, 138, 206, 10.1016/j.nlm.2016.10.010 Cornsweet, 1970 Cowansage, 2014, Direct reactivation of a coherent neocortical memory of context, Neuron, 84, 432, 10.1016/j.neuron.2014.09.022 Crick, 1984, Memory and molecular turnover, Nature, 312, 101, 10.1038/312101a0 D’Avila Garcez, 2015 Denny, 2014, Hippocampal memory traces are differentially modulated by experience, time, and adult neurogenesis, Neuron, 83, 189, 10.1016/j.neuron.2014.05.018 Edmiston, 2017, Visual interference disrupts visual knowledge, Journal of Memory and Language, 92, 281, 10.1016/j.jml.2016.07.002 Elbert, 1997, Input-increase and input-decrease types of cortical reorganization after upper extremity amputation in humans, Experimental Brain Research, 117, 161, 10.1007/s002210050210 Enoki, 2009, Expression of long-term plasticity at individual synapses in hippocampus is graded, bidirectional, and mainly presynaptic: Optical quantal analysis, Neuron, 62, 242, 10.1016/j.neuron.2009.02.026 Fauth, 2019, Self-organized reactivation maintains and reinforces memories despite synaptic turnover, eLife, 8, e43717, 10.7554/eLife.43717 Fernandez-Rodriguez, 2015, Memory and combinatorial logic based on DNA inversions: Dynamics and evolutionary stability, ACS Synthetic Biology, 10.1021/acssynbio.5b00170 Florence, 1998, Large-scale sprouting of cortical connections after peripheral injury in adult macaque monkeys, Science, 282, 1117, 10.1126/science.282.5391.1117 Fodor, 1975 Franke, 2016, Structures of neural correlation and how they favor coding, Neuron, 89, 409, 10.1016/j.neuron.2015.12.037 Freas, 2018, Landmark learning, cue conflict, and outbound view sequence in navigating desert ants, Journal of Experimental Psychology: Animal Learning and Cognition, 44, 409 Frisch, 1965 Furber, 2016, Large-scale neuromorphic computing systems, Journal of Neural Engineering, 13, 051001, 10.1088/1741-2560/13/5/051001 Gallistel, 1990 Gallistel, 2017, Finding numbers in the brain, Phil. Trans. R. Soc. B, 373, 20170119, 10.1098/rstb.2017.0119 Gallistel, 2017, The neurobiological bases for the computational theory of mind, 275 Gallistel, 2017, The coding question, Trends in Cognitive Science, 21, 498, 10.1016/j.tics.2017.04.012 Gallistel, 2014, Time to rethink the neural mechanisms of learning and memory, Neurobiology of Learning and Memory, 108, 136, 10.1016/j.nlm.2013.11.019 Gallistel, 2019, Contingency, contiguity, and causality in conditioning: Applying information theory and Weber’s Law to the assignment of credit problem., Psychological Review, 126, 761, 10.1037/rev0000163 Gallistel, 2010 Gallistel, 2013, The neuroscience of learning: Beyond the Hebbian synapse, Annual Review of Psychology, 64, 169, 10.1146/annurev-psych-113011-143807 Gao, 2018, Persistent increases of PKMζ in sensorimotor cortex maintain procedural long-term memory storage, iScience, 5, 90, 10.1016/j.isci.2018.07.002 Garson, 1993, Must we solve the binding problem in neural hardware?, Behavioral and Brain Sciences, 16, 459, 10.1017/S0140525X00031022 Gehring, 1998 Gerstner, 2014 Ghandour, 2019, Orchestrated ensemble activities constitute a hippocampal memory engram, Nature Communications, 10, 2637, 10.1038/s41467-019-10683-2 Grant, 2019, The synaptomic theory of behavior and brain disease, Cold Spring Harbor Symposia on Quantitative Biology, 037887 Groh, 2001, Converting neural signals from place codes to rate codes, Biological Cybernetics, 85, 159, 10.1007/s004220100249 Halder, 1995, Induction of ectopic eyes by targeted expression of the eyeless gene in Drosophila, Science, 267, 1788, 10.1126/science.7892602 Hayashi-Takagi, 2015, Labelling and optical erasure of synaptic memory traces in the motor cortex, Nature, 525, 333, 10.1038/nature15257 Hebb, 1949 Hecht-Nielsen, 2012 Heo, 2018, Identification of long-lived synaptic proteins by proteomic analysis of synaptosome protein turnover, Proceedings of the National Academy of Sciences of the United States of America, 115, E3827, 10.1073/pnas.1720956115 Hofer, 2009, Experience leaves a lasting structural trace in cortical circuits, Nature, 457, 313, 10.1038/nature07487 Horner, 2015, Evidence for holistic episodic recollection via hippocampal pattern completion, Nature Communication, 6, 7462, 10.1038/ncomms8462 Hu, 2007, Emotion enhances learning via norepinephrine regulation of AMPA-receptor trafficking, Cell, 131, 160, 10.1016/j.cell.2007.09.017 Huang, 1998, Synaptic plasticity: Going through phases with LTP, Current Biology, 8, R350, 10.1016/S0960-9822(98)70219-2 Hubel, 1959, Receptive fields of single neurones in the cat’s striate cortex, The Journal of Physiology, 148, 574, 10.1113/jphysiol.1959.sp006308 Huber, 2015, Egocentric and geocentric navigation during extremely long foraging paths of desert ants, Journal of Comparative Physiology A, 10.1007/s00359-015-0998-3 Isaac, 1995, Evidence for silent synapses: Implications for the expression of LTP, Neuron, 15, 427, 10.1016/0896-6273(95)90046-2 Jackson, 2013, Recall of spatial patterns stored in a hippocampal slice by long-term potentiation, Journal of Neurophysiology, 110, 2511, 10.1152/jn.00533.2013 Jirenhed, 2007, Acquisition, extinction, and reacquisition of a cerebellar cortical memory trace, The Journal of Neuroscience, 27, 2493, 10.1523/JNEUROSCI.4202-06.2007 Jirenhed, 2011, Time course of classically conditioned Purkinje cell response is determined by initial part of conditioned stimulus, Journal of Neuroscience, 31, 9070, 10.1523/JNEUROSCI.1653-11.2011 Jirenhed, 2011, Learning stimulus intervals—adaptive timing of conditioned Purkinje cell responses, The Cerebellum, 10, 523, 10.1007/s12311-011-0264-3 Jirenhed, 2016, Are Purkinje cell pauses drivers of classically conditioned blink responses?, The Cerebellum, 15, 10.1007/s12311-015-0722-4 Jirenhed, 2017, Learned response sequences in cerebellar Purkinje cells, Proceedings of the National Academy of Sciences of the United States of America, 114, 6127, 10.1073/pnas.1621132114 Johansson, 2015, Activation of a temporal memory in purkinje cells by the mGluR7 receptor, Cell Reports, 13, 1741, 10.1016/j.celrep.2015.10.047 Johansson, 2014, Memory trace and timing mechanism localized to cerebellar Purkinje cells, Proceedings of the National Academy of Science of the United States of America, 111, 14930, 10.1073/pnas.1415371111 Johansson, 2018, Absence of parallel fibre to Purkinje cell LTD during eyeblink conditioning, Scientific Reports, 8, 14777, 10.1038/s41598-018-32791-7 Johansson, F. (2015). A Purkinje cell timing mechanism: On the physical basis of a temporal duration memory. (PhD), Lund, Lund, Sweden. Johnson, 2009, Hebbian reverberations in emotional memory micro circuits, Frontiers in Neuroscience, 3, 198, 10.3389/neuro.01.027.2009 Josselyn, 2015, Finding the engram, Nature Reviews Neuroscience, 16, 521, 10.1038/nrn4000 Juusola, 2007, Coding with spike shapes and graded potentials in cortical networks, Bioessays, 29, 178, 10.1002/bies.20532 Kiela, 2011 Kim, 2017, Encoding of discriminative fear memory by input-specific LTP in the amygdala, Neuron, 95, 1129, 10.1016/j.neuron.2017.08.004 Kiya, 2011, Dance type and flight parameters are associated with different mushroom body neural activities in worker honeybee brains, PLoS One, 6, e19301, 10.1371/journal.pone.0019301 Koch, 1997, Computation and the single neuron, Nature, 385, 207, 10.1038/385207a0 Koch, 1999 Kyrke-Smith, 2018, Bridging synaptic and epigenetic maintenance mechanisms of the engram, Frontiers in Molecular Neuroscience, 11, 369, 10.3389/fnmol.2018.00369 Lacagnina, 2019, Distinct hippocampal engrams control extinction and relapse of fear memory, Nature Neuroscience, 22, 753, 10.1038/s41593-019-0361-z Langille, 2018, The synaptic theory of memory: A historical survey and reconciliation of recent opposition, Frontiers in Systems Neuroscience, 12, 52, 10.3389/fnsys.2018.00052 Lansner, 2003, Cell assembly dynamics in detailed and abstract attractor models of cortical associative memory, Theory in Biosciences, 122, 19, 10.1078/1431-7613-00072 Lashley, 1950, In search of the engram, 454 Laughlin, 2001, Energy as a constraint on the coding and processing of sensory information, Current Opinion in Neurobiology, 11, 475, 10.1016/S0959-4388(00)00237-3 Laughlin, 2004, The implications of metabolic energy requirements for the representation of information in neurons, 187 Laughlin, 1998, The metabolic cost of neural information, Nature Neuroscience, 1, 36, 10.1038/236 Leibovic, 1990 Levy, 1979, Synapses as associative memory elements in the hippocampal formation, Brain Research, 175, 233, 10.1016/0006-8993(79)91003-5 Lisman, 1985, A mechanism for memory storage insensitive to molecular turnover: A bistable autophosphorylating kinase, Proceedings of the National Academy of Sciences of the United States of America, 82, 3055, 10.1073/pnas.82.9.3055 Lisman, 2018, Memory formation depends on both synapse-specific modifications of synaptic strength and cell-specific increases in excitability, Nature Neuroscience, 21, 309, 10.1038/s41593-018-0076-6 Lisman, 2013, The theta-gamma neural code, Neuron, 77, 1002, 10.1016/j.neuron.2013.03.007 Liu, 2012, Optogenetic stimulation of a hippocampal engram activates fear memory recall, Nature, 484, 381, 10.1038/nature11028 Lundqvist, 2013, Attractor hypothesis of associative cortex: Insights from a biophysically detailed network model, functional brain mapping and the endeavor to understand the working brain, Francesco Signorelli and Domenico Chirchiglia, IntechOpen, Lundstrom, 2006, Decoding stimulus variance from a distributional neural code of interspike intervals, The Journal of Neuroscience, 26, 9030, 10.1523/JNEUROSCI.0225-06.2006 Ma, 2016, Re-silencing of silent synapses unmasks anti-relapse effects of environmental enrichment, Proceedings of the National Academy of Sciences of the United States of America, 113, 5089, 10.1073/pnas.1524739113 McClelland, 1989, Parallel distributed processing: Implications for cognition and development, 9 McClelland, 1986, The appeal of parallel distributed processing, 365 McCulloch, 1943, A logical calculus of the ideas immanent in nervous activity, Bulletin of Mathematical Biophysics, 7, 115, 10.1007/BF02478259 Menzel, 2011, A common frame of reference for learned and communicated vectors in honeybee navigation, Current Biology, 21, 645, 10.1016/j.cub.2011.02.039 Miller, 2016, Itinerancy between attractors states in neural systems, Current Opinions in Neurobiology, 40, 14, 10.1016/j.conb.2016.05.005 Miller, 2007, Covalent modification of DNA regulates memory formation, Neuron, 53, 857, 10.1016/j.neuron.2007.02.022 Miota, 2004, Putting fear in its place: Remapping of hippocampal place cells during fear conditioning, The Journal of Neuroscience, 24, 7015, 10.1523/JNEUROSCI.5492-03.2004 Mongillo, 2017, Intrinsic volatility of synaptic connections – a challenge to the synaptic trace theory of memory, Current Opinions Neurobiology, 46, 7, 10.1016/j.conb.2017.06.006 Moore, 2019, Piwi/PRG-1 argonaute and TGF-β mediate transgenerational learned pathogenic avoidance, Cell, 177, 1827, 10.1016/j.cell.2019.05.024 Mozzachiodi, 2008, Changes in neuronal excitability serve as a mechanism of long-term memory for operant conditioning, Nature Neuroscience, 11, 1146, 10.1038/nn.2184 Murphy, 2004, Increased neuronal excitability, synaptic plasticity, and learning in aged Kvβ1.1 knockout mice, Current Biology, 14, 1907, 10.1016/j.cub.2004.10.021 Navabi, 2014, Engineering a memory with LTD and LTP, Nature, 511, 348, 10.1038/nature13294 Newell, 1958, Elements of a theory of human problem solving, Psychological Review, 65, 151, 10.1037/h0048495 Nicoll, 2005, Synaptic plasticity at hippocampal mossy fibre synapses, Nature Reviews Neuroscience, 6, 863, 10.1038/nrn1786 O’Connor, 2005, Graded bidirectional synaptic plasticity is composed of switch-like unitary events, Proceedings of the National Academy of Sciences of the United States of America, 102, 9679, 10.1073/pnas.0502332102 O’Reilly, 2006, Biologically based computational models of high-level cognition, Science, 314, 91, 10.1126/science.1127242 Panzeri, 2015, Neural population coding: Combining insights from microscopic and mass signals, Trends in Cognitive Sciences, 19, 162, 10.1016/j.tics.2015.01.002 Panzeri, 2001, The role of spike timing in the coding of stimulus location in rat somatosensory cortex, Neuron, 29, 769, 10.1016/S0896-6273(01)00251-3 Park, 2011, Ensemble place codes in hippocampus: CA1, CA3 and dentate gyrus place cells have multiple place fields in large environments, PLoS One, 6, e22349, 10.1371/journal.pone.0022349 Park, 2016, Neuronal allocation to a hippocampal engram, Neuropsychopharmacology, 41, 2987, 10.1038/npp.2016.73 Petersen, 1998, All-or-none potentiation at CA3-CA1 synapses, Proceedings of the National Academy of Sciences of the United States of America, 95, 4732, 10.1073/pnas.95.8.4732 Petrovic, 2017, Metabotropic action of postsynaptic kainite receptors triggers hippocampal long-term potentiation, Nature Neuroscience, 20, 529, 10.1038/nn.4505 Pignatelli, 2019, Engram cell excitability state determines the efficacy of memory retrieval, Neuron, 101, 274, 10.1016/j.neuron.2018.11.029 Poo, 2016, What is memory? The present state of the engram, BMC Biology, 14, 40, 10.1186/s12915-016-0261-6 Posner, 2019, Neuronal small RNAs control behavior transgenerationally, Cell, 177, 1814, 10.1016/j.cell.2019.04.029 Putnam, 1975, The nature of mental states, Mind, language, and reality: philosophical papers, Vol. 2, 429, 10.1017/CBO9780511625251.023 Quiring, 1994, Homology of the eyeless gene of the Drosophila to the small eye gene in mice and aniridia in humans, Science, 265, 785, 10.1126/science.7914031 Quiroga, 2012, Concept cells: The building blocks of declarative memory functions, Nature Reviews Neuroscience, 13, 587, 10.1038/nrn3251 Rao-Ruiz, 2019, Engram-specific transcriptome profiling of contextual memory consolidation, Nature Communications, 10, 2232, 10.1038/s41467-019-09960-x Recanzone, 2000, Frequency and intensity response properties of single neurons in the auditory cortex of the behaving macaque monkey, Journal of Neurophysiology, 83, 2315, 10.1152/jn.2000.83.4.2315 Riley, 2005, The flight paths of honeybees recruited by the waggle dance, Nature, 435, 205, 10.1038/nature03526 Roe, 1992, Visual projections routed to the auditory pathway in ferrets: Receptive fields of visual neurons in primary auditory cortex, The Journal of Neuroscience, 12, 3651, 10.1523/JNEUROSCI.12-09-03651.1992 Romo, 2003, Flutter discrimination: Neural codes, perception, memory and decision making, Nature Reviews Neuroscience, 4, 203, 10.1038/nrn1058 Roy, 2016, Memory retrieval by activating engram cells in mouse models of early Alzheimer's disease, Nature, 531, 508, 10.1038/nature17172 Roy, 2017, Silent memory engrams as the basis for retrograde amnesia, Proceedings of the National Academy of Sciences of the United States of America, 114, E9972, 10.1073/pnas.1714248114 Ryan, 2015, Engram cells retain memory under retrograde amnesia, Science, 348, 1007, 10.1126/science.aaa5542 Salz, 2016, Time cells in hippocampal area CA3, The Journal of Neuroscience, 36, 7476, 10.1523/JNEUROSCI.0087-16.2016 Samsonovich, 1997, Path integration and cognitive mapping in a continuous attractor neural network model, The Journal of Neuroscience, 17, 5900, 10.1523/JNEUROSCI.17-15-05900.1997 Sanders, 2012, Elimination of dendritic spines with long-term memory is specific to active circuits, The Journal of Neuroscience, 32, 12570, 10.1523/JNEUROSCI.1131-12.2012 Saucier, 1995, Spatial learning without NMDA receptor-dependent long-term potentiation, Nature, 378, 186, 10.1038/378186a0 Scudellari, 2015, Inner Workings: DNA for data storage and computing, Proceedings of the National Academy of Sciences of the United States of America, 112, 15771, 10.1073/pnas.1520100112 Sehgal, 2018, Memory allocation mechanisms underlie memory linking across time, Neurobiology of Learning and Memory, 153, 21, 10.1016/j.nlm.2018.02.021 Shannon, 1948, A mathematical theory of communication, Bell Systems Technical Journal, 27, 623, 10.1002/j.1538-7305.1948.tb00917.x Silva, 2009, Molecular and cellular approaches to memory allocation in neural circuits, Science, 326, 391, 10.1126/science.1174519 Sloman, 1978 Smolensky, 1986, Information processing in dynamical systems: Foundations of harmony theory, Vol. 1, 194 Stevens, 2018, Conserved features of the primate face code, Proceedings of the National Academy of Sciences of the United States of America, 115, 584, 10.1073/pnas.1716341115 Tanaka, 2014, Cortical representations are reinstated by the hippocampus during memory retrieval, Neuron, 84, 347, 10.1016/j.neuron.2014.09.037 Tang, 2018, Recurrent computations for visual pattern completion, Proceedings of the National Academy of Sciences of the United States of America, 115, 8835, 10.1073/pnas.1719397115 Teyler, 1986, The hippocampal memory indexing theory, Behavioral Neuroscience, 100, 147, 10.1037/0735-7044.100.2.147 Teyler, 2007, The hippocampal indexing theory and episodic memory: Updating the index, Hippocampus, 17, 1158, 10.1002/hipo.20350 Thiele, 2017, Neuromodulation of attention, Neuron, 97, 769, 10.1016/j.neuron.2018.01.008 Tonegawa, 2015, Memory engram storage and retrieval, Current Opinion in Neurobiology, 35, 101, 10.1016/j.conb.2015.07.009 Trettenbrein, 2016, The demise of the synapse as the locus of memory: A looming paradigm shift, Frontiers in Systems Neuroscience, 10, 88, 10.3389/fnsys.2016.00088 Turing, 1947, Lecture on the automatic computing engine Turing, 1950, Computing machinery and intelligence, Mind, 59, 433, 10.1093/mind/LIX.236.433 Ungerleider, 2011, Uncovering the visual “alphabet”: Advances in our understanding of object perception, Vision Research, 51, 782, 10.1016/j.visres.2010.10.002 Viosca, 2009, Enhanced CREB-dependent gene expression increases the excitability of neurons in the basal amygdala and primes the consolidation of contextual and cued fear memory, Learning & Memory, 16, 193, 10.1101/lm.1254209 von Neumann, 1958 von Neumann, J. (1945/2011). First draft of a report on the EDVAC, with an Introduction by Michael D. Godfrey. Electrical Engineering. Moore School of Engineering. Philadelphia. https://fa82ee93-a-62cb3a1a-s-sites.googlegroups.com/site/michaeldgodfrey/vonneumann/vnedvac.pdf?attachauth=ANoY7coIXpfrfxAc6WIumKsVMDFLrSTyPXfyN0reBvTKXI0p_n3wD1cjYCVy6nnhUBAx8Yu386kEWKZutq627_U9mgYflcVF4WTndPAg0DP0fINIl_aEhZ3nGuhFIECTLZ5VAyoXbOi-1KxoUeZ0pPgwHrtPVAV-dwEV7Vps_OViS72YWgVa9pfe57yhAYWIxOoFYOFxev7GFu7i8-QFXMvolRnIFV2y5mIRgAQ-kLDTjLkudVKrhks%3D&attredirects=0. Watrous, 2015, Phase-amplitude coupling supports phase coding in human ECoG, eLife, 4, e07886, 10.7554/eLife.07886 Wetmore, 2014, Bidirectional plasticity of purkinje cells matches temporal features of learning, The Journal of Neuroscience, 34, 1731, 10.1523/JNEUROSCI.2883-13.2014 Whitlock, 2006, Learning induces long-term potentiation in the hippocampus, Science, 313, 1093, 10.1126/science.1128134 Wilkes, J. T., and Gallistel, C. R. (2016). Information theory, memory, prediction, and timing in associative learning (original long version). https://osf.io/preprints/y9sg2/. Wilkes, 2017, Information theory, memory, prediction, and timing in associative learning, 481 Wittlinger, 2006, The ant odometer: Stepping on stilts and stumps, Science, 312, 1965, 10.1126/science.1126912 Wittlinger, 2007, The desert ant odometer: A stride integrator that accounts for stride length and walking speed, Journal of Experimental Biology, 210, 198, 10.1242/jeb.02657 Wolfe, 2012, The binding problem lives on: Comment on De Lillo, Trends in Cognitive Science, 16, 307, 10.1016/j.tics.2012.04.013 Yang, 2009, Stably maintained dendritic spines are associated with lifelong memories, Nature, 462, 920, 10.1038/nature08577 Yassa, 2013, Competitive trace theory: A role for the hippocampus in contextual interference during retrieval, Frontiers in Behavioral Neuroscience, 7, 107, 10.3389/fnbeh.2013.00107 Yiu, 2014, Neurons are recruited to a memory trace based on relative neuronal excitability immediately before training, Neuron, 83, 722, 10.1016/j.neuron.2014.07.017 Zeldenrust, 2018, Neural coding with bursts—current state and future perspectives, Frontiers Computational Neuroscience, 12, 48, 10.3389/fncom.2018.00048 Zhang, 2003, The other side of the engram: Experience-driven changes in neuronal intrinsic excitability, Nature Reviews Neuroscience, 4, 885, 10.1038/nrn1248 Zhou, 2009, CREB regulates excitability and the allocation of memory to subsets of neurons in the amygdala, Nature Neuroscience, 12, 1438, 10.1038/nn.2405 Zhu, 2018, Architecture of the mouse brain synaptome, Neuron, 99, 781, 10.1016/j.neuron.2018.07.007