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SYSTEMOGENY Function: Traditional and Systemic Instead of regarding, e.g.,views “vision”, “audition”, “attention”, “motor control” and “motivation” as functions, a fundamentally different definition of a “function” was elaborated in functional system theory. In this theory, function refers to an achievement of a result through the realization of a certain functional system. Such a function cannot be attributed not only to certain neural networks but even to the brain. It relates to the whole organism. Systemogenesis versus Organogenesis Systemogeney and Ontogeny During early ontogeny, those differently localized elements undergo selective and accelerated maturation that is essential for achieving the results of the systems, providing for the survival of an organism at the early stages of individual development. • Systemogeny takes place not only during the early ontogenetic period, but also during adult development • The formation of a new behavioural act is always a formation of a new system The essence of the basic idea underlying the absolute majority of studies within the framework of the ABC paradigm A sso cia tion ism B e h aviorism C on n ectio n ism A B C le arning (D .C .D e n ne tt) A n early illustration of a reflex , from D escartes' L'hom m e (1664) Cartesian concept of animal spirits "The traces exist because those pores in the brain that a spirit passed through before are more likely to open when it approaches the again. And then the spirit can pass through these pores more easily." Long-Term Potentiation and Evoked Spike Responses in the Cingulate Cortex of Freely Mobile Rats A. G. Gorkin, K. G. Reymann, and Yu. I. Aleksandrov Neuroscience and Behavioral Physiology, Vol. 33, No. 8, 2003 Long-term potentiation of EP in the cingulate cortex after tetanic stimulation of the subiculo-cingulate tract. The ordinate shows the EP rise rate, % of initial, and the abscissa shows time, h. The time during which posttetanic EP was significantly (t test, p < 0.05) different from initial is noted by the line and asterisks. Re-juvenilization Nowadays it is commonly accepted that many regularities of modification of functional and morphologic characteristics of neurones, as well as of control of gene expression, serve as a basis for the formation of adaptive behaviour in adults, and are comparable to those found at the early ontogenetic stages. Early genesLate genesMorphological modifications Neuronal death that follows seizures, cerebral ischemia, etc is initiated by overactivation of glutamate receptors, and an increased intracellular concentration of Ca2+. Increased concentration of Ca2+ causes transcriptional activiation of so-called “cell-death genes”. Immediate-early genes play a central role in converting extracellular signals into long-lasting alterations in cellular phenotype. The existing data provide strong evidence for a relationship between PERSISTENT activation of immediate-early genes and delayed neuronal death (that is a form of programmed cell death, or apopotosis). A similar relationship between persistent expression of c-fos, and cell death in neural and non-neural systems was apparent. By contrast, TRANSIENT induction of immediate-early genes seems to occur in neuronal populations that are destined to survive. SS. Schreiber & M. Baudry Trends Neurosci., 1995, 18, 446. It can be suggested that the expression of c-Fos is a prerequisite that allows other transcription components carry out their specific decision for death or survival. (p.2736) Lee Y., Park K.H., Baik S.H., Cha Ch.I. Attenuation of c-Fos basal expression in the cerebral cortex of aged rat. NeuroReport, 1998, 9, 2733-2736 Compensation NEONEUROGENESIS ? Primary and/or Secondary assortment? Pathological state ?? LG EG Secondary assortment or Systemogenesis Systemic specialization Normal state Learning Adults Early onotogeny Maturation Primary assortment Systemic pre-specialization death ? DG EG EG Neuron Neuron is always active Each step in neuronal elimination is active. Neurons of different classes express receptors whose activation can lead to their death. C. Raoul, Pettmann B. & Henderson C.E. Active killing of neurons during development and following stress. Current Opin. In Neurobiol. 2000, 10, 111-117 Altruism and Suicide Altruism and social cheating in the social amoeba Dictyostelium discoideum. Nature, 2000, 408, 965 Strassmann J.E., Zhu Y. & Queller D.C. Most of the time, D. discoideum lives as haploid, free-living, amoeboid cells that divide asexually. When starved, 104-105 of these cells aggregate into a slug. The anterior 20% of the slug altruistically differentiates into a non-viable stalk, supporting the remaining cells, most of which become viable spores. If aggregating cells come from multiple clones, there should be selection for clones to exploit other clones by contributing less than their proportional share to the sterile stalk. Half of the chimaeric mixtures show a clear cheater and victim. Thus, unlike the clonal and highly cooperative development of most multicellular organisms, the development of D. discoideum is partly competitive, with conflicts of interests among cells. Four deaths and a funeral: from caspases to alternative mechanisms. Nature Reviews, 2001, 2, 1-10 Leist M. & Jaattela M. Cell suicide in the adult nervous system has serious implications for the whole organism Selection instead of Instruction Both in biology and in cognitive sciences there is a principled discontinuation of the search for a meager stock of all-purpose innate “primitives”, out of which everything else ought to be “constructed” through a process of “enrichment”, mediated by “association” … The present trend is to grant a very rich innate repertoire and then look for the mechanisms of internal selection. The key role of the environment is not that of supplying more structure, but rather that of steering a selection. We should conceive of “learning” as a process of triggering, filtering, parameter-setting, progressive specialization and selection. A SELECTIVE THEORY IS BY NOW ALWAYS THE MOST PLAUSIBLE CANDIDATE FOR EVERY PROCESS OF ACQUISITION [Piattelli-Palmarini M. Evolution, selection and cognition: From “learning” to parameter setting in biology and in the study of language. Cognition, 1989, 31, 1-44] The system-selective concept of learning “Place-cells” in rats “Face neurons” in monkeys “Face neurons” in human beings «Gnostic neurons» by Konorsky New Neurons Studies of information storage in model neural networks have shown that adding new learning sequentially to the network can result in "catastrophic interference" [McCloskey M. & Cohen N.J., 1989]. New information can obscure previously stored information. If new memories got new neurons with new connections, catastrophic interference might be avoided. (W.T.Greenough et al., Nature Neurosci., 1999) Studies in our laboratory supported the suggestion that the mechanism of the formation of behaviour (learning) is the recruitment (specialization) of “new” (previously silent) neurons into the subserving of specific new behaviours. Data from other laboratories (Jog et al. 1999, Thompson & Best 1990, Wilson & McNaughton 1993, Swadlow & Hicks1997, Woodward et al. 1998) confirms the suggestion that learning involves “new” neurons rather than "re-learning" (the replacement of preexisting specialization by a new one) of the “old” cells; that newly formed unit specialization remains the same during the whole period of the recording (weeks and even months), and that there are many silent neurons in different brain areas that may become active at some time. Immune Cells Never Forget Once learned, some abilities, such as swimming or riding a bike, are never forgotten even after years without practice. Others, say running a marathon, need a regular brushing up. Immunologists have long debated which category our immunological memory falls into. Once immune cells learn to recognize a particular antigen, such as a viral protein, do they need constant reminders to stay on top of things, or are their memories permanent? Two reports in this week's issue of Science … now bolster the notion that immune cells never forget. … Most experts are convinced that memory T cells don't need constant stimulation … to stay in shape Michael Hagmann Science vol. 286, # 5444, 1999: 1266 - 1267 Konorski’s Concept of Gnostic Areas and Units Unitary perceptions are mediated by the discharge of single neurons, called gnostic units. Gnoctic areas are considered as files of gnostic units representing all the unitary perceptions established in a given subject. Once a potential gnostic unit had been preemted by a particular stimulus pattern so as to become transformed into an actual gnostic unit representing that unitary perception, it become resistant to any new stimulus pattern. [Konorski J. Integrative activity of the brain. An interdisciplinary approach. Univ. Chicago Press: Chicago, 1967, 531 p.] BEHAVIORAL SPECIALIZATIONS OF NEURONS Retrosplenial cortex DISTRIBUTION OF FOSFOS-POSITIVE NEURONS Motor cortex 25 * 10% 3 Retrosplenial cortex * 20 Motor cortex 3 56% 53% 1 37% 2 5% Fos-positive cells 1 2 (% of total in the areas) 39% 15 10 5 0 - unidentified neurons - non-specifically task-related neurons - specifically task-related neurons O «C L» RO NT T SI UI CQ «A IO N» T N- I O UIS «N CQ A IO N» «M E NT AI NC NA E» *p<0.05 versus the motor cortex. Intensity of c-Fos expression is the highest in the retrosplenial cortex which has a higher number of specific behaviorally specialized neurons RELATIOSHIPS BETWEEN c-FOS EXPRESSION AND BEHAVIORAL SPECIALIZATION OF NEURONS Retrosplenial cortex 100 m 0.23% bar-pressing related neurons 23.6% Motor cortex 100 m 0.04% 3.6% 0.98% out of Fos-positive 1.11% NeoNeurogenesis Newly generated neurons in the adult are not only affected by the formation of a hippocamapl-dependent memory, but also participate in it. The new cells are about 1-2 weeks of age when they become involved in the learned response. This ability to undergo rapid structural change may be a characteristic of immature neurons that makes them ideally suited for forming associations between stimuli. Newly generated neurons may not be used for learning under more lenient conditions, but become involved as task demands increase. [T.J. Shors, G. Miesegaes, A. Beylin, M. Zhao, T. Rydel, and Gould E. Neurogenesis in the adult is involved in the formation of trace memories. Nature, 2001, 410, 372-375] NEW SYSTEMS Behavioral Act 2 OLD SYSTEMS Individual development Behavioral Act 1 The new system is added to the existing systems; newly formed systems do not substitute for previously existing systems, but are "superimposed" over them. What does it mean - "to superimpose, but not to substitute"? Many experiments have demonstrated that a complex instrumental behaviour is mastered not only through the realization of new systems, that were formed during the process of learning the acts comprising the behaviour, but also by the simultaneous realization of older systems, that had been formed at previous stages of individual development. 3 2 REAR WALL 3 2 1 FRONT WALL 4 4 1 8 6 7 AP PP 5 AF LIF FS 8 7 FG LIF 6 RC 5 5 0 1.0 1.0 .8 .8 .6 .6 .4 .4 .2 .2 Behavioral Act 2 NEW SYSTEMS Behavioral Act 1 OLD SYSTEMS Individual development 1 sec 1 2 3 4 5 6 7 8 9 10 1 2 3 4 5 6 7 8 9 10 OLD SYSTEMS Individual development Behavioral Act 1 NEW SYSTEMS Behavioral Act 2 OLD SYSTEMS Individual development NEW SYSTEMS Behavioral Act 1 Behavioral Act 2 500 msec Example of the activity of the M-neuron that was activated during rightward body movement. Activations of this neuron appear during the turning to the right when approaching the pedal on one side of the cage (FC), side of the cage (RC) or during this movement in defensive behavior (DB). NEW SYSTEMS Behavioral Act 2 OLD SYSTEMS Individual development Behavioral Act 1 The realization of behaviour is the realization of the history of behavioural development, that is, of many systems, each fixing a certain stage of development of the given behaviour. Behavior is a simultaneous realization of a set of system ranging from the most archaic to the newest ones Hippocampus Position EMG Actogr. rear NEW SYSTEMS Behavioral Act 1Behavioral Act 2 Neuron Neuron Actogr. front Actogr. rear Neuron Actogr. front Actogr. rear OLD SYSTEMS Motor cortex from a hand from the floor Trigeminal mesencephalic nucleus Neuron Jaw mov. EMG 500 msec Individual development Actogr. front Memory Environmental afferentation Parameters of the result Acceptor of action’s result Decision making Triggering stimulus Environmental afferentation Program of the action Result of the action Action Motivation Afferent synthesis r 1 r R n 2 Act n r 1 T r 2 Act n+1 r R n + 1 3 r 1 T r 2 Act n+2 r 3 RE -CONSOLIDATION the modifications of preexisting memory Definitive behavior mismatch MEMORY Re-consolidation in consequence of memory reactivation LEARNING Behavioral Act 2 • SPECIALIZATION - the recruitment of new neurones into the subserving of specific new behaviours • ACCOMMODATIVE RE CONSOLIDATION - the modifications of pre-existing behaviours neurones NEW SYSTEMS Behavioral Act 1 OLD SYSTEMS Individual development TWO CLOSELY CONNECTED PROCESSES: Memory “projection” to cerebral structures Patterns of neurons’ behavioral specialization in different brain structures % 100 NEW 60 40 20 0 LimbC Hpc VisC S-sC Hpt MotC OptT Act 1 Act 2 OLD SYSTEMS UNIDENTIFIED NEW SYSTEMS OLD Individual development 80 Our recordings of unit activity in many cerebral structures during instrumental behavior have demonstrated that neurons with new specializations are abundant in cerebral cortex, whereas phylogenetically archaic and peripheral structures contain very few of them. E- E+ 28% Acute ethanol 11% Chronic ethanol HEALTHY 34% 17% L>M 22% ALCOHOLIC 34% M>L M>L Acute ethanol 30% 26% L~M The specificity of memory “projection” to cerebral structures is determined by the characteristics of neurons composing these structures. These characteristics determine the involvement of neurons of the given structure into the formation of the certain behavior. Brain Research 876 (2000) 154–165 Effects of ethanol on hippocampal place-cell and interneuron activity * Aaron M. White , Phillip J. Best Definitive behavior Mismatch MEMORY Re-consolidation in consequence of memory reactivation At the beginning of learning Pre-exiting memory 1-EGE Mismatch 2-Tets Mismatch Pre-exiting memory 3-Success Mismatch Pre-exiting memory 4-«Altruistic suicide» Pre-exiting memory 5-Consolidation of a new memory Pre-exiting memory 6-Modification of pre-exiting memory Pre-exiting memory New memory Accommodative re-consolidation 7-Dynamics of memory Pre-exiting memory Re-consolidation New in consequence of memory memory reactivation COGNITIVE SOLIPSISM Cognitive science still proceeds as if culture did not matter. The central assumption of cognitive solipsism is that the mind may be, indeed must be, conceptualized as a system that is contained entirely inside a box. In the case of vertebrates like ourselves, that box happens to be the brain. The strong form of this doctrine holds that [mind] basic structure is a biological given, structured according to a set of innate neuropsychological universals. Within this framework, culture is necessarily assigned a secondary role. Lev Vygotsky … was one of the first to recognize the symbiosis of the developing mind with culture… [Merlin Donald The central role of culture in cognitive evolution: a reflection on the myth of the “isolated mind”. In: Culture, thought, and development. L.P.Nucci, G. Saxe, E. Turiel (Eds.) Lawrence Erlbaum Associates, Inc., 2000, pg. 19-38] THE UNIT OF CULTURE We recommended that the unit of culture – now called meme – be the same as the node of semantic memory and its correlates in brain activity “Nodes”, or reference points, in semantic memory … can be associated with neural activity in the brain Nodes are almost always linked to other nodes, so that to recall one node is to summon others The linkage of nodes is assembled as a hierarchy to organize information with more and more meaning [Edward O. Wilson. Consilience. The unity of knowledge. A.A. Knoff. New York. 1998, pp. 135,136] Human specializations and brain activity Bird and car experts were tested with functional magnetic resonance imaging during tasks with … cars and birds. The right fusiform girus and occipital lobe showed significant expertise effect. [I. Gauthier et al. Expertise for cars and birds recruits brain areas involved in face recognition. Nature Neurosci. 2000 3 (2), 191 –197] THE IMPLICATIONS OF NICHE CONSTRUCTION FOR EVOLUTION Standard Evolutionary Theory Extended Evolutionary Theory Laland et al. 1996, 1999, 2001; Odling-Smee et al. 2003 THE IMPLICATIONS OF NICHE CONSTRUCTION FOR HUMAN SCIENCES Cultural niche construction can influence not only subsequent cultural changes, but also some human genetic changes Gene-culture coevolution he cultural trait of dairy farming is thought to have evolved in association with the gene for lactose absorption