Generation of variants of a motor act in a modular and hierarchical motor network

BACKGROUND: Most motor systems can generate a variety of behaviors, including categorically different behaviors and variants of a single motor act within the same behavioral category. Previous work indicated that many pattern-generating interneuronal networks may have a modular organization and that distinct categories of behaviors can be generated through flexible combinations of a small number of modules or building blocks. However, it is unclear whether and how a small number of modules could possibly generate a large number of variants of one behavior. RESULTS: We show that the modular feeding motor network of Aplysia mediates variations in protraction duration in biting-like programs. Two descending commands are active during biting behavior and trigger biting-like responses in a semiintact preparation. In the isolated CNS, when activated alone, the two commands produce biting-like programs of either long or short protraction duration by acting specifically on two modules that have opposite effects on protraction duration. More importantly, when coactivated at different frequencies, the two commands produce biting programs with an intermediate protraction duration. CONCLUSIONS: It was previously hypothesized that behavioral variants may be produced by combining different activity levels of multiple descending commands. Our data provide direct evidence for such a scheme and show how it is implemented in a modularly organized network. Thus, within a modular and hierarchical architecture, in addition to generating different categories of behavior, a small number of modules also efficiently implements variants of a single behavior.     

Relation between reaction time and the characteristics of a motor act

The connection of reaction time (RT) with spatial-temporal motor parameters was studied in humans. Duration of the motor act was set by experimenter. In response to the signal the tested person pressed one or two buttons according to instruction. The interval between these pressings corresponded to the time of performing the movement. It is shown, that RT significantly depends on the regime of work, duration of movement, and distance between buttons: with greater distances such dependence becomes significant. It is suggested that dependence of RT upon various motor act parameters is determined by differences in the levels of brain structures activation and in spatial-temporal organization of the movement.     

The structural organization and neurochemical mechanisms of the participation of the nucleus accumbens in the interaction of the limbic and motor systems and in the regulation of motor behavior]

The review of own and literature data devoted to structural and neurochemical organization of the nucleus accumbens (Acc) as well as spatial organization of their projection fibers in comparison with nucleus caudatus (neostriatum) has been presented. The facts concerned with correlations revealed between both the cell clusters and histochemical compartments as well as the compartmental organization of afferent and efferent striatal connections were analyzed. The presented data propose the existence of sensorimotor and limbic parts of the dorsal and ventral striatum, which are involved in the parallelly functioning systems. The common and different signs of these two systems and its role in the regulation of the movement behaviour has been proposed. A lot of attention also was payed to the Acc and the neostriatum interaction. The many pathways by which Acc can influence neostriatum functions and therefore the motor activity controlled by the neostriatum are discussed. It was shown that the Acc can influence on the striatal synaptic DA release. The sign of this influence depended upon DA/glutamic acid interactions in the Acc. It was stressed that the influence of Acc on striatal DA-ergic system is very likely mediated via kainate/quisqualate (but not NMDA) inputs of the neostriatum. The balance of DA-ergic mechanisms of neostriatum and Acc as important basis of animals adequate behaviour in conditioning situation was proposed. The disbalance of these mechanisms could leads to pathology.     

The stepping motor protein as a feedback control ratchet

It is explained how going from a one headed motor protein to a stepping two headed motor protein is equivalent to going from a stochastically flashing ratchet to a feedback control ratchet. Both these ratchets have been well studied in the literature and their speeds and efficiencies are briefly reviewed. Next it is shown how a feedback control ratchet mechanism model can account for very accurate recent data obtained on kinesin. Finally, the role of internal friction in the operation of stepping motor proteins is discussed.     

Interhemispheric motor asymmetry of conditioned feedback

The interhemispheric asymmetry and lateralization of motor functions, as applied to backward conditioning, was studied in two dogs that afterwards turned to be a right-hander and a left-hander. In the process of food-procuring instrumental conditioning it was revealed that one of the dogs predominantly used the right paw (with the dominance of the left hemisphere) and the second animal used the left paw (the right brain hemisphere dominated). The earliest signs of lateralization of the motor functions were observed in the backward conditioned connections (which had been formed earlier than the direct connections or direct conditioned reflexes). Consequently, the presence and sign of asymmetry can be predicted on the basis of manifestation of the backward connections observed at the early stages of acquisition of food-procuring behavior. The backward connections are strictly specific and have the same lateralization as the direct ones.     

Reducing auditory feedback and the acquisition of motor response timing

Three groups of subjects (n=10) attempted to move a lever 50 cm along a track in 1.50 sec under one of three auditory feedback conditions: Fully augmented increasing auditory feedback (FAF) in which a constant level of velocity-related auditory feedback was provided for all 25 learning trials, reducing auditory feedback (FAF) in which the level of feedback was progressively reduced over the learning trials, and no auditory feedback (NAF). All subjects performed 10 trials with no auditory feedback after a 10-min rest interval. The hypothesis that acquisition of the criterion task would be facilitated under RAF compared to FAF derived partial support. It was concluded that there is sufficient evidence to justify further investigation of reducing auditory feedback as a technique of motor skill acquisition.     

Cell adhesion and migration in the organization of spinal motor neurons

Spinal motor neurons are critical to the ability of animals to move and thus essential to survival. Motor neurons that project axons to distinct limb-muscle targets are topographically organized such that central nervous system position reflects the location of the muscle in the limb. The central positioning of limb-projecting motor neurons arises during development through motor neuron migration followed by a period of coalescence into discrete groupings of motor neurons which project axons to an individual muscle. These so-called motor pools are a common feature of motor organization in higher vertebrates. Recent work has highlighted the critical role for armadillo family member catenin-dependent functions of the cadherin family of cell adhesion molecules in directing the organization of motor neurons. Cadherin function appears to be important for both the motor neuron migration and coalescence phases of the emergence of motor neuron topography. Here, I review this recent work in the context of our understanding of the general development of spinal motor neurons.     

Possible organization pattern of interaction between the generators of cyclic motor activity and inputs from supraspinal and afferent systems

Possible organization patterns of scratching and locomotor generators that allow interpretation of experimentally demonstrated reorganizations in temporal parameters of these generator activities after electrical stimulation of descending and peripheral afferent systems were analyzed with application of mathematical simulation of neuronal generator systems. The results obtained led to the conclusion that patterns of such reorganizations influenced by signals from suprasegmental and/or peripheral systems may be determined by only two factors: 1) the structure of synaptic connections between interneuronal functional groups underlying these generator associations, and 2) the structure of connections between these groups of interneurons and fibers from suprasegmental and peripheral afferent sources. The existence of inhibitory-excitatory actions from descending and afferent systems upon the neurons of locomotor or scratching generator half-centers is a sufficient condition to ensure phasic changes in the sensitivity of these generators to supraspinal and afferent signals. The locomotor generator, unlike the scratching generator, is apparently characterized by a more complex organization of connections between functional neuronal groupings and descending fibers.Translated from Neirofiziologiya, Vol. 25, No. 1, pp. 45–50, January–February, 1993.     

Cell adhesion and migration in the organization of spinal motor neurons

Spinal motor neurons are critical to the ability of animals to move and thus essential to survival. Motor neurons that project axons to distinct limb-muscle targets are topographically organized such that central nervous system position reflects the location of the muscle in the limb. The central positioning of limb-projecting motor neurons arises during development through motor neuron migration followed by a period of coalescence into discrete groupings of motor neurons which project axons to an individual muscle. These so-called motor pools are a common feature of motor organization in higher vertebrates. Recent work has highlighted the critical role for armadillo family member catenin-dependent functions of the cadherin family of cell adhesion molecules in directing the organization of motor neurons. Cadherin function appears to be important for both the motor neuron migration and coalescence phases of the emergence of motor neuron topography. Here, I review this recent work in the context of our understanding of the general development of spinal motor neurons.     

Traveling-wave pattern generator controls movement and organization of sensory feedback in a spinal cord model

 A traveling wave in a two-dimensional spinal cord model constitutes a stable pattern generator for quadruped gaits. In the context of the somatotopic organization of the spinal cord, this pattern generator is sufficient to generate stable locomotive limb trajectories. The elastic properties of muscles alone, providing linear negative feedback, are sufficient to stabilize stance and locomotion in the presence of perturbative forces. We further show that such a pattern generator is capable of organizing sensory processing in the spinal cord. A single-layer perceptron was trained to associate the sensory feedback from the limb (coding force, length, and change of length for each muscle) with the two-dimensional activity profile of the traveling wave. This resulted in a well-defined spatial organization of the connections within the spinal network along a rostrocaudal axis. The spinal network driven by peripheral afferents alone supported autonomous locomotion in the positive feedback mode, whereas in the negative feedback mode stance was stabilized in response to perturbations. Systematic variation of a parameter representing the effect of gamma-motor neurons on muscle spindle activity in our model led to a corresponding shift of limb position during stance and locomotion, resulting in a systematic displacement alteration of foot positions. Received: 30 July 2001 / Accepted in revised form: 17 April 2002 Correspondence to: A. Kaske (e-mails: alexander.kaske@mtc.ki.se, alexander.kaske@vglab.com)     

Microtubule motor proteins and the organization of the pollen tube cytoplasm

Molecular motors are molecules that drive a wide range of activities (for example, organelle movement, chromosome segregation, and flagellar movement) in cells. Thus, they play essential roles in diverse cellular functions. Understanding their structures, mechanisms of action and different roles is therefore of great practical importance. The role of microtubules during pollen tube growth is presently not identified, nor are basic properties. We do not know, for example, where microtubules are organized, the extent of microtubule dynamics, and the polarity of microtubules in the pollen tube. Roles of microtubules and related motors in organelle trafficking are not clear. Regardless of scarce information, microtubule-based motors of both the kinesin and dynein families have been identified in the pollen tube. Most of these microtubule motors have also been found in association with membrane-bounded organelles, which suggest that these proteins could translocate organelles or vesicles along microtubules. The biochemical features of these proteins are typical of the motor protein class. Immunofluorescence microscopy of pollen tubes probed with antibodies that cross-react with microtubule motors indicate that these proteins are localized in different regions of the pollen tube; therefore, they could have different roles. Although a number of microtubule motors have been identified in the pollen tube, the role of these proteins during pollen tube germination and growth or organelle movement is not yet recognized, as tube elongation and organelle movement in the pollen tube depend mostly on actin filaments. In the effort to understand the specific role that microtubules and related motors have in the pollen tube, it is therefore necessary to identify the molecular machinery that interacts with microtubules. Furthermore, it is crucial to clearly establish the types of interaction between organelles and microtubules. This review summarizes the current state of the art on microtubule motors in the pollen tube, mainly surrounding the putative roles of microtubule motors in organelle movement and cytoplasmic organization. Some hypotheses and speculations are also presented.     

Chaos in multi-looped negative feedback systems

Non-linear control systems with multiple negative feedback loops display periodicity, quasiperiodicity and period-doubling bifurcations leading to chaos. The possibility that normal fluctuations in physiological control may result from deterministic chaos in multi-looped negative feedback systems is discussed.     

On the stability of equilibria in metabolic feedback systems

A sufficient criterion for the stability of the equilibrium of a metabolic feedback system will be constructed.     

Synchronization of motor units and its simulation in parallel feedback system

Comparison of the discharge patterns of soleus motor units with associated changes in force exerted by foot during quiet stance have already demonstrated the following facts. In the initial stage of standing, the motor units exhibited stationary and asynchronous discharges. The force showed a sporadic presence of the high frequency oscillation in the 8–10 Hz band. After five to ten minutes of standing, the firing rate of individual motor unit discharges increased to about 10 spikes/sec and discharges of each motor unit were synchronized and phase-locked to each of the accompanying force oscillation. During this transitional stage, the discharges of motor units were characterized by spike dropouts from an otherwise regular spike train. To simulate the changes in the discharge characteristics, we have proposed a parallel feedback model of the stretch reflex arc. This was made of multiple -motoneurons, motor units and muscle spindles. And motor units interact each other through group Ia afferent signals. As a result of simulation, motor units were found to exhibit stationary and asynchronous discharges when feedback gain was kept small. With an increase of feedback gain, the firing rate of individual motor units increased and finally the discharges of them were synchronized. During this transitional stage, the spike dropouts were observed in accordance with the experimental results. The neuronal mechanism of synchronization may partly be explained by the interactions of motor neurons through the above stated parallel feedback system.     

Bifurcation and chaotic behaviour in simple feedback systems

Simple feedback systems are well understood. For example recent work in control theory has provided methods for predicting the existence and stability of periodic solutions in the output of such systems. These methods do not consider the possibility that behaviour more complicated than simple limit cycles might occur. Further, the control engineering literature does not cite either practical examples, or theoretical analyses, of systems behaving chaotically.In this paper we show that chaotic behaviour can occur in simple feedback systems of finite dimensions, and compare such systems with a family of discrete maps studied by May.