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Cutaneous afferents are responsible for a wide range of sensations discount sildigra 100mg mastercard, but most are also capable of modula- Withdrawal responses ting motor behaviour through spinal best sildigra 50 mg, supraspinal Theseresponseshaveaspinalpathwayandarecom- and transcortical pathways cheap sildigra 100 mg on line. There is a tendency for monly but erroneously thought to involve a flexor clinicians to group all cutaneous afferents together purchase sildigra 120mg overnight delivery, synergy activated by a nociceptive stimulus buy sildigra 50 mg visa. With- and this creates confusion, leads to the usage of dif- drawal reflexes have a specific organisation, are rea- ferent terms for the same function and the same sonablystereotyped,andareelicitedbyconvergence term for different functions, and makes the systems of noxious and tactile stimuli (cf. A thesis of this flexion reflex, the corresponding pathways were book, addressed in many chapters, is that cutaneous named FRA (flexor reflex afferent) pathways. The nociceptive withdrawal vant pathways is inhibited by activation of pathways (flexion) reflex was subsequently shown to be poly- mediating short-latency FRA reflexes. The organisa- synaptic (for references, see Hunt & Perl, 1960), and tionoflong-latencyFRApathwayssuggeststhatthey this was confirmed by intracellular recordings from play a role in the generation of locomotor stepping motoneurones (R. Eccles & Lundberg, 1959;Holmqvist & The above responses can be generated by stimu- Lundberg, 1961). Further investigations showed that lating cutaneous afferents in isolation. In addition, administration of DOPA in the acute spinal cat cutaneous afferents contribute to shaping the motor suppressed short-latency FRA responses, releasing output through their extensive convergence on transmission in a long-latency FRA pathway, which interneurones interposed in pathways fed by mus- had a half-centre organisation, capable of gener- cleafferentsorcorticospinalvolleys(cf. Chapters3–7 ating alternating activation of extensors and flexors and 10), and onto PAD interneurones mediat- (Jankowska et al. However, when cutaneous and FRA volleys elicit different effects in Initial findings the same motoneurone(s), there is evidence for a specialised cutaneous pathway. Investigations of spinal reflexes received impetus from the work of Sherrington (1906, 1910)onthe Reflexes elicited by low-threshold nociceptive flexion reflex. He showed that, in the cutaneous afferents spinalised decerebrate animal, noxious skin stim- The toe extensor reflex of the cat uli excite flexors and inhibit extensors in the ipsilat- eral hindlimb (the flexion reflex), accompanied by This is the most clear-cut example of a specialised excitation of extensors and inhibition of flexors in cutaneous reflex. Gentle pressure on the central the contralateral limb (the crossed extension reflex). The ensuing movement gives rise to an impulse flow in FRA which is channelled back into the reflex already activated, so that its activity is reinforced and prolonged (see p. From data in Engberg (1964)(a), and modified from Schouenborg (2002)(b), Lundberg (1973, 1979)((c), (d)), and Baldissera, Hultborn & Illert (1981)(e), with permission. Background from animal experiments 387 response in the plantar flexors of the toes (i. This reflex is due ingmotoneurones,presumablydesignedfordiscrete to the activation of slowly adapting mechanorecep- movementsofthedifferentdigits(Sasakietal. This extensor activation Cutaneousreflexesduringlocomotionarealsomedi- is appropriate to avoid the stimulus. Inchronicspinal specific relationship between receptive field, acti- cats walking on a treadmill, tactile stimuli applied to vated muscle(s) and the resulting reflex withdrawal thedorsumofthepawevokeshort-latencyresponses has been revealed both in the rat and the cat (for involving the flexors during the swing phase, but review, see Schouenborg, 2002). The responses in ate cutaneous receptive field corresponding to the knee muscles are stronger and have shorter laten- skin area withdrawn by contraction of the partic- cies than those of ankle and hip muscles. This is illustrated for the recep- ternandtimingofactivation(i)distinguishtheabove tive fields for withdrawal reflexes involving the pero- responses from FRA-induced responses, which are neus longus, tibialis anterior and biceps of the cat in stereotyped and synchronous in all flexors, and (ii) Fig. Isolatedkneeflexion (ii) Nociceptors and, to a lesser extent, slowly early in swing is sufficient to overcome the obstacle adapting low-threshold mechanoreceptors provide touched by the pad dorsum, whereas the increased the afferent input to withdrawal reflex pathways. Projections to motoneurones innervating Reflexes in the forelimb slow- and fast-twitch motor units In the forelimb of the cat, low-threshold cutaneous A differentiation between FRA pathways and spe- afferents project to segmental interneurones inter- cialised cutaneous pathways has also been pos- posed in proprioceptive pathways (cf. Introduction, sible in the motoneurones innervating fast-twitch and Hongo et al. Alstermark & Lundberg, 1992; Chapter are excited from cutaneous afferents but inhibited 10,p. Burke, Jankowska & evokes highly specialised reflexes in digit motoneu- ten Bruggencate, 1970). A descending action on specialised reflex pathways from skin has been inferred because facilitation of cutaneous effects may occur without concomi- FRA reflex pathways tant changes in the FRA effects evoked from high- threshold muscle afferents. Flexor reflex afferents (FRA) (i) Rubrospinal facilitation of low-threshold cuta- neous excitation of extensor motoneurones inner- FRA include group III and, in the cat, group II vatingfastmotorunits(cf. Because of this convergence, above for specialised cutaneous pathways and in descending excitation of the relevant interneurones Chapter 7 for group II pathways. There are two may receive feedback reinforcement by impulses reflex patterns from the FRA: the short-latency evokedfromtheplantarcushionduringcontactwith (early) reflexes found in the acute spinal cat, and the ground (see Fig.

The hierarchical organization of biological structural units from the cellular to the organismal levels (cell organelles cheap 120mg sildigra, nuclei buy sildigra 25mg without a prescription, neurons discount 100 mg sildigra with amex, synapses discount sildigra 50mg line, neural groups sildigra 100mg low cost, nervous tissue, and cerebral organs), naturally suggested a hierarchical representa- tion of a system. However, the hierarchical aspect of the corresponding functional organization is far from evident. The novel three-dimensional representation of a biological system that one of us has proposed (G. Chauvet, 1996a), with axes for space scales, time scales, and structural units, allows visualization of the coupling between the structural and functional organizations. This representation is based es- sentially on the determination of the time scales of the dynamic systems describing physiological functions. This functional hierarchy is useful for determining the phys- iological functions associated with nervous structures. In the case of real neural net- works, there are at least two physiological functions: the propagation of membrane potential on a time scale on the order of milliseconds, and the modification of synap- tic e‰cacy on a time scale on the order of seconds or even hours. Thus, the func- tional order has its origin in a functional hierarchy that is evidently a manifestation of molecular mechanisms. Typically, the artificial neural networks generally studied have several neuron layers. The structure-function rela- tionship is more evident in this representation than in any other one. The hierarchical network is fundamentally di¤erent and, in particular, possesses specific emergent properties, that is, properties that appear at a higher level in a new structure. An im- Inputs Outputs (behaviors) Sensory Hierarchical Motor neurons circuit neurons Figure 7. Properties emerge from a lower level and appear at a higher level inside a new structure. This new structure is called a functional unit if, and only if, it has a specific function. Mathematical Modeling of Neuromimetic Circuits 133 portant advantage of the hierarchical representation is that it o¤ers a rigorous ap- proach to the notion of a functional unit that may now be defined as a structural unit with a specific function at a higher level of organization (G. The functional unit, possessing its own time scale, incorporates a new function that can be derived mathematically from the lower levels of organization in a biological system. For example, a neuromimetic circuit may be considered as a functional unit. Hierarchical Representation of a Biological Theory of Functional Organization Functional Interactions In the course of our work on physiological models, ranging from the molecular to the organismal levels (G. Chauvet, 1996b), some novel ideas specific to the study of biology have been introduced, in particular the concepts of nonsymmetric and non- local functional interactions in hierarchical space. These basic concepts emerged from a bottom-up approach to living systems; that is, from a systematic study of iso- lated physiological functions, followed by the integration of these functions at the level of the organism. A significant consequence of this theory is that living organ- isms can be given not only a double organizational representation that is simultane- ously structural and functional but also a double mathematical representation that is simultaneously geometric and topological. We may compare it to a mathematical function in the sense that the action of one structure on another results in a certain product. The physiological function would then be the action (the application, in mathematical terms) and the product would be the result of the function (the value of the function, in mathematical terms) that is often identified with the physiological function itself. Although this definition is general, it is unfortunately not operational. It is relatively easy to describe particular physiological functions such as vision, di- gestion, memorization, and so on, but it is far more di‰cult to give an operational definition of a physiological function in general. One possibility may be to define a physiological function in terms of a combinatorial set of functional interactions between structures. Such functional interactions are evidently specific since they de- scribe the action (whatever its nature) of one structure on another or, more precisely, the action of a source on a sink, after the action has undergone a transformation in the source. In addition, it has another important property, that of nonlocality, a notion somewhat more di‰- cult to appreciate since it stems from the structural hierarchy of the system (G. This may be explained as follows: (1) From a mathematical point of view, in a continuous representation, the action of one structure on another is necessarily the 134 G. This does not correspond to the action of one cell on another in physical space since a cell contains regions with specialized functions and therefore cannot be reduced to a point.

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