Principle of neuronal organization of defensive reflexes in mollusks

Spontaneous and evoked synaptic activity of command neurons for the defensive response of spiracle closing were studied by simultaneous intracellular recording of activity of several identified CNS neurons in snails. Comparison of monosynaptic EPSPs in command neurons evoked by discharges of presynaptic neurons with spontaneous synaptic potentials indicated that the central organization of the defensive reflex is in the form of a two-layered neuron net in which each neuron of the afferent layer possesses a local receptive field, but which overlaps with other afferent neurons. Each neuron of the afferent layer is connected with each neuron of the efferent layer by monosynaptic excitatory connections that differ in efficiency (maximal only with one neuron of the efferent layer). Both receptive fields of neurons of the afferent layer and "fields of efficiency of synaptic connections" are distributed according to the normal law. As a result of this organization the neuron net acquires a new quality: The action of different stimuli leads to the appearance of differently located "spatial excitation profiles" of efferent layer neurons even when this action of the stimulus occurs not at the center of the receptive field.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 16, No. 1, pp. 26–34, January-February, 1984.     

A neural network model for selective attention in visual pattern recognition

A neural network model of the mechanism of selective attention in visual pattern recognition is proposed and simulated on a digital computer.When a complex figure consisting of two patterns or more is presented to the model, it is segmented into individual patterns, and each pattern is recognized separately. Even if one of the patterns to which the model is paying selective attention is affected by noise or defects, the model can recall the complete pattern from which the noise has been eliminated and the defects corrected. It is not necessary for perfect recall that the stimulus pattern should be identical in shape to the training pattern. Even though the pattern is distorted in shape or changed in size, it can be correctly recognized and the missing portions restored.The model consists of a hierarchical neural network which has efferent as well as afferent connections between cells. The afferent and the efferent signals interact with each other in the network: the efferent signals, that is, the signals for selective attention, have a facilitating effect on the afferent ones, and, at the same time, the afferent signals gate efferent signal flow. When some feature in the stimulus is not extracted in the afferent paths, the threshold for detection of that feature is automatically lowered by decreasing the efficiency of inhibition, and the model tries to extract even vague traces of the undetected feature.     

Consciousness and Reflex

An analogy is drawn between the functions of consciousness and the model of a reflex. Consciousness consists of an afferent part, which is the perception of the external world; a central part, which is thought; and an efferent part, which is a decision concerning an action and the transmission of a motor command. At the basis of higher mental functions lies the unification of reflexes into complexes; and within these complexes, key structures are differentiated that synthesize qualitatively disparate information. The centers of integration for the effectuation of the afferent and efferent functions of consciousness are located in the posterior and anterior sections of the brain, respectively. Predominantly afferent and efferent stages, with localized foci of interaction, may also be distinguished in thought. The communicative function of consciousness is realized primarily by centers in the left hemisphere. A discrepancy between the afferent and efferent functions of consciousness may underlie certain forms of mental pathology.     

Lymphocyte traffic through granulomas: Differences in the recovery of indium-111-labeled lymphocytes in afferent and efferent lymph

Afferent lymphatics draining granulomas and efferent lymphatics from normal and stimulated lymph nodes were cannulated in sheep. There was a greatly increased output of cells in afferent lymph-draining chronic inflammatory sites or Freund's adjuvant-induced granulomas. Cells collected from these lymphatics were radiolabeled with ¹¹¹In and injected intravenously. The reappearance of these labeled cells in lymph at various sites was measured. Labeled afferent lymph cells migrated from blood through the granuloma back into afferent lymph in large numbers and with kinetics which were comparable to efferent lymphocytes recirculating through a lymph node. When labeled afferent lymph cells were injected the specific activity (cpm/10⁷ cells) in afferent lymph was five times higher than that in efferent lymph from a normal node. When efferent lymph cells were labeled the afferent lymph specific activity was one-half that in efferent lymph. It is suggested that the cells in afferent lymph migrate preferentially from blood through the granuloma and constitute a unique population of cells.     

Efferent fibers innervate gustatory and mechanosensitive afferent fibers in frog fungiform papillae

A possibility of efferent innervation of gustatory and mechanosensitive afferent fiber endings was studied in frog fungiform papillae with a suction electrode. The amplitude of antidromic impulses in a papillary afferent fiber induced by antidromically stimulating an afferent fiber of glossopharyngeal nerve (GPN) with low voltage pulses was inhibited for 40 s after the parasympathetic efferent fibers of GPN were stimulated orthodromically with high voltage pulses at 30 Hz for 10 s. This implies that electrical positivity of the outer surface of papillary afferent membrane was reduced by the efferent fiber-induced excitatory postsynaptic potential. The inhibition of afferent responses in the papillae was blocked by substance P receptor blocker, L-703,606, indicating that substance P is probably released from the efferent fiber terminals. Slow negative synaptic potential, which corresponded to a slow depolarizing synaptic potential, was extracellularly induced in papillary afferent terminals for 45 s by stimulating the parasympathetic efferent fibers of GPN with high voltage pulses at 30 Hz for 10 s. This synaptic potential was also blocked by L-703,606. These data indicate that papillary afferent fiber endings are innervated by parasympathetic efferent fibers.     

Consciousness and reflex

In the paper an analogy is drawn between functions of consciousness and scheme of reflex. Consciousness includes afferent part perception of external world, central one--thinking, and efferent one--decision about action and motor command presentation. The basis of higher psychic functions is the junction of reflexes in complicated complexes, singling out in their composition key structures realizing synthesis of qualitatively different information. The centers of integration at realization of afferent functions of consciousness are situated in posterior, and at efferent ones--in anterior brain areas. In the process of thinking predominantly afferent and efferent stages may also be singled out with corresponding localization of foci of interaction. Communicative function of consciousness is provided mainly by centers of the left hemisphere. Breakage between afferent and efferent functions of consciousness may be the basis of some forms of psychic pathology as depression.     

Renorenal reflexes: interaction between efferent and afferent renal nerve activity.

In rats, stimulation of renal mechanoreceptors by increasing ureteral pressure results in a contralateral inhibitory renorenal reflex response consisting of increases in ipsilateral afferent renal nerve activity, decreases in contralateral efferent renal nerve activity, and increases in contralateral urine flow rate and urinary sodium excretion. Mean arterial pressure is unchanged. To study possible functional central interaction among the afferent renal nerves and the aortic and carotid sinus nerves, the responses to renal mechanoreceptor stimulation were compared in sinoaortic denervated rats and sham-denervated rats before and after vagotomy. In contrast to sham-denervated rats, there was an increase in mean arterial pressure in response to renal mechanoreceptor stimulation in sinoaortic-denervated rats. However, there were no differences in the renorenal reflex responses among the groups. Thus, our data failed to support a functional central interaction among the renal, carotid sinus, and aortic afferent nerves in the renorenal reflex response to renal mechanoreceptor stimulation. Studies to examine peripheral interaction between efferent and afferent renal nerves showed that marked reduction in efferent renal nerve activity produced by spinal cord section at T6, ganglionic blockade, volume expansion, or stretch of the junction of superior vena cava and right atrium abolished the responses in afferent renal nerve activity and contralateral renal function to renal mechanoreceptor stimulation. Conversely, increases in efferent renal nerve activity caused by thermal cutaneous stimulation increased basal afferent renal nerve activity and its responses to renal mechanoreceptor stimulation. These data suggest a facilitatory role of efferent renal nerves on renal sensory receptors.     

Cardiogenic reflexes after immune injury of the heart

Afferent and efferent spike activity from the parasympathetic (vagus) and sympathetic cardiac nerves were recorded simultaneously with ECG, and indices of heart function were measured in acute experiments on anesthetized dogs, which allowed us to study the modifications of cardio-cardiac reflex influences after a local immune heart injury. After an injury nidus has been formed in the heart, cardiogenic depressor reflexes evoked by an intracoronary application of veratrine or bradykinin were considerably suppressed or even abolished, and afferent spike activity in the vagus cardiac nerves noticeably decreased. At the same time, both the facilitation of activity in sympathetic afferent fibers and pressor reflex effects were preserved after the heart injury. Different localization of vagus and sympathetic afferent structures in the heart and their specialized sensitivity to the biologically active substances are suggested as the factors determining the pattern of cardiogenic reflex influences after a heart injury.Neirofiziologiya/Neurophysiology, Vol. 27, No. 1, pp. 18–25, January–February, 1995.     

Suppression of antigraft immunity by preimmunization. III. Characterization of suppressor T cells involved in suppression of the efferent phase of DTH against alloantigens

Suppressor T (Ts) cells that can suppress delayed type hypersensitivity (DTH) against histocompatibility (H) antigens can be isolated from spleen and lymph nodes a few days after i.v. immunization of mice with irradiated allogeneic spleen cells. In this paper we investigated the suppression of the efferent phase of DTH to characterize the Ts cells involved, and to compare them with the afferent phase Ts cells that have been characterized in a previous paper of this series. The DTH against third party alloantigens that were not used for the i.v. suppressive immunization could be suppressed by presenting the third party alloantigens together with the original alloantigens in the challenge inoculum for eliciting the DTH reaction. Thus the ultimate suppressive effect by the Ts cells that are active during the efferent phase of DTH is nonspecific. This non-specific suppression of DTH to alloantigens has previously been found for the afferent phase Ts cells as well. For suppression of the efferent phase of DTH to alloantigens, a population of Lyt-1+2+ Ts cells appeared to be essential, just like in the suppression of the afferent phase of DTH to alloantigens. We did not find evidence for the involvement of cyclophosphamide-sensitive auxiliary Ts cells in suppression of the efferent phase of DTH. Also no evidence was found for H-2 or Igh-restricted activation and function of the Ts cells that were active during afferent and efferent phases of the DTH response to H antigens. In view of these similarities between afferent phase and efferent phase Ts cells we conclude that there are no arguments as yet to suppose that there is more than one type of T cells involved in the suppression of the afferent and efferent limb of DTH against H antigens.     

Populations of behavior-reactive neurons in the monkey neostriatum

Comparative analysis of the unit activity of the monkey putamen during multistage behavior showed that neurons of the putamen are active during all the behavioral actions. It was established that the number of the behavior-related neurons changes considerably less than number of neurons which reorganize their activity at the time. Reorganization of unit activity in the putamen is considered as reflecting the efferent code which controls behavior, and the degree of reorganization--as a measure of change of this code in relation to organization of ongoing behavioral action. It has been discovered that the change in the number of the active neurons at various steps of behavior and reorganization of their activity occurs independently. It may be related to two main afferent systems of striatum: ascending from rhe brain stem, and corticofugal which brings differentiated information to the neuronal net of striatum from various parts of the cortex.     

Vascular anatomy of the gills of the stingarees Urolophus mucosus and U. paucimaculatus (Urolophidae,Elasmobranchii)

The branchial vascular anatomy of Urolophus mucosus and U. paucimaculatus was studied by scanning electron microscopical examination of critical-point-dried tissue or of vascular corrosion casts. The vasculature could be divided into arterioarterial and arteriovenous pathways, which channel the flow of blood through the gills. The arterioarterial pathway consists of an afferent branchial artery which gives rise to afferent distributing arteries that run through the tissues of the interbranchial septum and supply the afferent filament arteries of several filaments. Afferent filament arteries open regularly into a corpus cavernosum in the core of the filament; unlike other elasmobranchs no septal corpora cavernosa are found. At the tip of the filament, channels of the corpus cavernosum connect to a channel which passes across the distal end of the filament from afferent to efferent side. This channel always connects to the afferent filament artery, and in many filaments it connects to the efferent filament artery as well. In addition, a vascular arcade connects all the afferent filament arteries along the entire length of each hemibranch. The filament corpus cavernosum supplies the secondary lamellae. The lamellae drain into efferent lamellar arterioles which in turn drain into the efferent filament artery and the efferent branchial artery. The vascular anatomy of the arteriovenous pathway is similar to that described in other elasmobranchs and consists of arteriovenous anastomoses, found only arising from efferent arterial circulation, and the venolymphatic system, which is composed of the central venous sinus and the companion vessels.     

Afferent and efferent components of joint position sense; interpretation of kinaesthetic illusion

The present model of joint angle perception is based on the following hypotheses: the perception and control of joint angle are closely interrelated processes; central motor commands are adequately expressed by shifts of an equilibrium point resulting from the interaction of antagonistic muscles and a load; two fundamental commands-reciprocal (r) and coactivative (c) provide for changes in activity of the antagonistic muscle pair. The dependence of joint angle on static muscle torque and r and c commands is derived (Eq. 5). The following principles of joint position sense are formulated: 1) the r component of the efferent copy plays the role of a reference point which shifts during voluntary regulation of muscle state, but remains unchanged during any passive alterations of joint position; 2) muscle afferent signals deliver not absolute but relative information (i.e. measured relatively to the central reference point). These signals turn out to be related to active muscle torque; 3) the nervous system evaluates muscle afferent signals on the basis of a scale determined by the level of coactivation of the antagonistic muscles. Kinaesthetic illusions appear to be due to disruptions in perception of afferent and/or efferent components of position sense. The present model is consistent with all the variety of kinaesthetic illusions observed experimentally. A qualitative neurophysiological schema for joint angle perception is proposed involving efferent copy and information concerning muscle torque delivered by the tendon organ, muscle spindle, and perhaps, articular receptors. It is known that the cerebellum incorporates both afferent and efferent information concerning movement. One may presume that it plays an essential role in position sense.     

Analysis of afferent,central, and efferent components of the baroreceptor reflex in mice

Studies of genetically modified mice provide a powerful approach to investigate consequences of altered gene expression in physiological and pathological states. The goal of the present study was to characterize afferent, central, and efferent components of the baroreceptor reflex in anesthetized Webster 4 mice. Baroreflex and baroreceptor afferent functions were characterized by measuring changes in renal sympathetic nerve activity (RSNA) and aortic depressor nerve activity (ADNA) in response to nitroprusside- and phenylephrine-induced changes in arterial pressure. The data were fit to a sigmoidal logistic function curve. Baroreflex diastolic pressure threshold (P(th)), the pressure at 50% inhibition of RSNA (P(mid)), and baroreflex gain (maximum slope) averaged 74 +/- 5 mmHg, 101 +/- 3 mmHg, and 2.30 +/- 0.54%/mmHg, respectively (n = 6). The P(th), P(mid), and gain for the diastolic pressure-ADNA relation (baroreceptor afferents) were similar to that observed for the overall reflex averaging 79 +/- 9 mmHg, 101 +/- 4 mmHg, and 2.92 +/- 0.53%/mmHg, respectively (n = 5). The central nervous system mediation of the baroreflex and the chronotropic responsiveness of the heart to vagal efferent activity were independently assessed by recording responses to electrical stimulation of the left ADN and the peripheral end of the right vagus nerve, respectively. Both ADN and vagal efferent stimulation induced frequency-dependent decreases in heart rate and arterial pressure. The heart rate response to ADN stimulation was nearly abolished in mice anesthetized with pentobarbital sodium (n = 4) compared with mice anesthetized with ketamine-acepromazine (n = 4), whereas the response to vagal efferent stimulation was equivalent under both types of anesthesia. Application of these techniques to studies of genetically manipulated mice can be used to identify molecular mechanisms of baroreflex function and to localize altered function to afferent, central, or efferent sites.     

Reflex regulation of airway smooth muscle tone.

Autonomic nerves in most mammalian species mediate both contractions and relaxations of airway smooth muscle. Cholinergic-parasympathetic nerves mediate contractions, whereas adrenergic-sympathetic and/or noncholinergic parasympathetic nerves mediate relaxations. Sympathetic-adrenergic innervation of human airway smooth muscle is sparse or nonexistent based on histological analyses and plays little or no role in regulating airway caliber. Rather, in humans and in many other species, postganglionic noncholinergic parasympathetic nerves provide the only relaxant innervation of airway smooth muscle. These noncholinergic nerves are anatomically and physiologically distinct from the postganglionic cholinergic parasympathetic nerves and differentially regulated by reflexes. Although bronchopulmonary vagal afferent nerves provide the primary afferent input regulating airway autonomic nerve activity, extrapulmonary afferent nerves, both vagal and nonvagal, can also reflexively regulate autonomic tone in airway smooth muscle. Reflexes result in either an enhanced activity in one or more of the autonomic efferent pathways, or a withdrawal of baseline cholinergic tone. These parallel excitatory and inhibitory afferent and efferent pathways add complexity to autonomic control of airway caliber. Dysfunction or dysregulation of these afferent and efferent nerves likely contributes to the pathogenesis of obstructive airways diseases and may account for the pulmonary symptoms associated with extrapulmonary disorders, including gastroesophageal reflux disease, cardiovascular disease, and rhinosinusitis.     

Morphometric characteristics of neuromuscular spindles in hypertrophied skeletal muscle

Skeletal muscle hypertrophy in young male rats was found to be accompanied by adaptive changes in neuromuscular spindles. The changes consisted in connective capsule thickening, increased diameter of NMS and intrafusal muscle fibers, expanded afferent and efferent nerve terminals, increased microcirculatory bed capacity. The quantitative and qualitative shifts observed in NMS structure are morphologically equivalent to the rise in their functional potential, which forms the basis for the functional changes in conditions of increasing skeletal muscle hypertrophy.     

A hierarchical neural network model for associative memory

A hierarchical neural network model with feedback interconnections, which has the function of associative memory and the ability to recognize patterns, is proposed. The model consists of a hierarchical multi-layered network to which efferent connections are added, so as to make positive feedback loops in pairs with afferent connections. The cell-layer at the initial stage of the network is the input layer which receives the stimulus input and at the same time works as an output layer for associative recall. The deepest layer is the output layer for pattern-recognition. Pattern-recognition is performed hierarchically by integrating information by converging afferent paths in the network. For the purpose of associative recall, the integrated information is again distributed to lower-order cells by diverging efferent paths. These two operations progress simultaneously in the network. If a fragment of a training pattern is presented to the network which has completed its self-organization, the entire pattern will gradually be recalled in the initial layer. If a stimulus consisting of a number of training patterns superposed is presented, one pattern gradually becomes predominant in the recalled output after competition between the patterns, and the others disappear. At about the same time when the recalled pattern reaches a steady state in he initial layer, in the deepest layer of the network, a response is elicited from the cell corresponding to the category of the finally-recalled pattern. Once a steady state has been reached, the response of the network is automatically extinguished by inhibitory signals from a steadiness-detecting cell. If the same stimulus is still presented after inhibition, a response for another pattern, formerly suppressed, will now appear, because the cells of the network have adaptation characteristics which makes the same response unlikely to recur. Since inhibition occurs repeatedly, the superposed input patterns are recalled one by one in turn.     

Vascular anatomy of the fish gill

The fish gill is the most physiologically diversified vertebrate organ, and its vasculature the most intricate. Application of vascular corrosion techniques that couple high-fidelity resins, such as methyl methacrylate, with scanning electron microscopy yields three-dimensional replicas of the microcirculation that have fostered a better appreciate gill perfusion pathways. This is the focus of the present review. Three vascular networks can be identified within the gill filament. The arterioarterial (respiratory) pathway consists of the lamellae and afferent and efferent segments of the branchial and filamental arteries and lamellar arterioles. The body of the filament contains two post-lamellar pathways: the interlamellar and nutrient. The interlamellar system is an extensive ladder-like network of thin-walled, highly distensible vessels that traverses the filament between, and parallel to, the lamellae and continues around the afferent and efferent borders of the filament. Interlamellar vessels are supplied by short, narrow-bore feeder vessels from the medial wall of the efferent filamental artery. A myriad of narrow-bore, tortuous arterioles arise from the basal efferent filamental artery and efferent branchial artery and anastomose to form the nutrient circulation of the arch and filament. In the filament body, nutrient capillaries and interlamellar vessels are often closely associated, and the former may ultimately drain into the latter. Many of the anatomical characteristics of interlamellar vessels are strikingly similar to those of mammalian lymphatic capillaries, with the exception that interlamellar vessels are directly fed by arteriovenous-like anastomoses. It is likely that gill interlamellar and mammalian lymphatics are physiologically, if not embryologically, equivalent.     

Inhibition and efferent facilitation of sensory activity in the isolated labyrinth of the frog

The functioning modalities of the efferent system were analysed in the isolated frog labyrinth. The efferent synapses of the posterior canal were activated via an axon reflex by antidromic electrical shocks (10-200 Hz) applied for increasing times (250 ms-10 s) to the anterior-horizontal nerves. Either decrease (inhibition) or increase (facilitation) in the resting discharge rate were observed in the majority of the units examined. Inhibition and facilitation, however, are peculiar to any given unit since inhibition does not reverse to facilitation or vice-versa. This fact as well as the long response latency (not less than 10 ms) and the linear dependence of both effects on the stimulation frequency suggest that inhibition and facilitation are due to the repetitive activation of two different types of efferent fibres synapsing on the hair cells. The drastic modifications in the afferent synaptic discharge produced by full activation of the efferent system indicate that the static properties (response asymmetry) as well as the dynamic properties (response adaptation) of the mechanically driven afferent response can be substantially controlled by the central nervous system at the receptor level.     

Vagal modulation of intestinal afferent sensitivity to systemic LPS in the rat

The central nervous system modulates inflammation in the gastrointestinal tract via efferent vagal pathways. We hypothesized that these vagal efferents receive synaptic input from vagal afferents, representing an autonomic feedback mechanism. The consequence of this vagovagal reflex for afferent signal generation in response to LPS was examined in the present study. Different modifications of the vagal innervation or sham procedures were performed in anesthetized rats. Extracellular mesenteric afferent nerve discharge and systemic blood pressure were recorded in vivo before and after systemic administration of LPS (6 mg/kg iv). Mesenteric afferent nerve discharge increased dramatically following LPS, which was unchanged when vagal efferent traffic was eliminated by acute vagotomy. In chronically vagotomized animals, to eliminate both vagal afferent and efferent traffic, the increase in afferent firing 3.5 min after LPS was reduced to 3.2 +/- 2.5 impulses/s above baseline compared with 42.2 +/- 2.0 impulses/s in controls (P < 0.001). A similar effect was observed following perivagal capsaicin, which was used to eliminate vagal afferent traffic only. LPS also caused a transient hypotension (<10 min), a partial recovery, and then persistent hypertension that was exacerbated by all three procedures. Mechanosensitivity was increased 15 min following LPS but had recovered at 30 min in all subgroups except for the chronic vagotomy group. In conclusion, discharge in capsaicin-sensitive mesenteric vagal afferents is augmented following systemic LPS. This activity, through a vagovagal pathway, helps to attenuate the effects of septic shock. The persistent hypersensitivity to mechanical stimulation after chronic vagal denervation suggests that the vagus exerts a regulatory influence on spinal afferent sensitization following LPS.