Learning, memory and a respiratory central pattern generator

In an attempt to elucidate the causal mechanisms underlying learning and memory we have developed a model system, aerial respiration in the pond snail Lymnaea stagnalis. A three-neuron central pattern generator (CPG) whose sufficiency and necessity have been demonstrated mediates this behaviour. Aerial respiration, while an important homeostatic behaviour, is inhibited by the activation of the whole body withdrawal response that the animal uses to protect itself. We found that it was possible to operantly condition snails not to perform aerial respiration in a situation, a hypoxic environment, where aerial respiration should predominate. Operant conditioning was achieved by eliciting the pneumostome withdrawal response, part of the whole body withdrawal response, each time the animal attempted to open its pneumostome to breathe. Yoked control animals did not demonstrate an alteration in breathing behaviour. Subsequently we determined neural correlates of this associative behaviour and found that neuronal changes are distributed throughout the CPG. This preparation may afford us the opportunity to determine the casual neuronal changes that underlie learning and memory of associative conditioning.     

A holographic model for associative memory chains

Holographic brain models are well suited to describe specific brain functions. Central nervous systems and holographic systems both show parallel information processing and non-localized storage in common. To process information both systems use correlation functions suggesting to develop cybernetical brain models in terms of holography. Associative holographic storage is done with two simultaneously existing patterns. They may reconstruct each other mutually. Time-sequentially existing patterns are connected to associative chains, if every two succeeding patterns do exist within a common period of time in order to be stored in pairs. Read out (recall) of associative chains—reconstructing coupled patterns which didn't exist simultaneously—requires advanced holographic techniques. Three different methods are described and tested experimentally. The underlying principles are feedback mechanisms, nonlinearities of the storage material and tridimensional architecture of the voluminous recording medium. Those principles evidently occur in neural storage systems supporting analogous information processing in neural- and holographic systems.     

Evolution of Associative Learning in Chemical Networks

Organisms that can learn about their environment and modify their behaviour appropriately during their lifetime are more likely to survive and reproduce than organisms that do not. While associative learning – the ability to detect correlated features of the environment – has been studied extensively in nervous systems, where the underlying mechanisms are reasonably well understood, mechanisms within single cells that could allow associative learning have received little attention. Here, using in silico evolution of chemical networks, we show that there exists a diversity of remarkably simple and plausible chemical solutions to the associative learning problem, the simplest of which uses only one core chemical reaction. We then asked to what extent a linear combination of chemical concentrations in the network could approximate the ideal Bayesian posterior of an environment given the stimulus history so far? This Bayesian analysis revealed the ‘memory traces’ of the chemical network. The implication of this paper is that there is little reason to believe that a lack of suitable phenotypic variation would prevent associative learning from evolving in cell signalling, metabolic, gene regulatory, or a mixture of these networks in cells.     

Multiple reward signals in the brain

The fundamental biological importance of rewards has created an increasing interest in the neuronal processing of reward information. The suggestion that the mechanisms underlying drug addiction might involve natural reward systems has also stimulated interest. This article focuses on recent neurophysiological studies in primates that have revealed that neurons in a limited number of brain structures carry specific signals about past and future rewards. This research provides the first step towards an understanding of how rewards influence behaviour before they are received and how the brain might use reward information to control learning and goal-directed behaviour.     

Speed/accuracy trade-off between the habitual and the goal-directed processes

Instrumental responses are hypothesized to be of two kinds: habitual and goal-directed, mediated by the sensorimotor and the associative cortico-basal ganglia circuits, respectively. The existence of the two heterogeneous associative learning mechanisms can be hypothesized to arise from the comparative advantages that they have at different stages of learning. In this paper, we assume that the goal-directed system is behaviourally flexible, but slow in choice selection. The habitual system, in contrast, is fast in responding, but inflexible in adapting its behavioural strategy to new conditions. Based on these assumptions and using the computational theory of reinforcement learning, we propose a normative model for arbitration between the two processes that makes an approximately optimal balance between search-time and accuracy in decision making. Behaviourally, the model can explain experimental evidence on behavioural sensitivity to outcome at the early stages of learning, but insensitivity at the later stages. It also explains that when two choices with equal incentive values are available concurrently, the behaviour remains outcome-sensitive, even after extensive training. Moreover, the model can explain choice reaction time variations during the course of learning, as well as the experimental observation that as the number of choices increases, the reaction time also increases. Neurobiologically, by assuming that phasic and tonic activities of midbrain dopamine neurons carry the reward prediction error and the average reward signals used by the model, respectively, the model predicts that whereas phasic dopamine indirectly affects behaviour through reinforcing stimulus-response associations, tonic dopamine can directly affect behaviour through manipulating the competition between the habitual and the goal-directed systems and thus, affect reaction time.     

Problems of learning and memory: one or multiple memory systems?

Learning, and hence memory, is ubiquitous not only throughout the animal kingdom, but apparently throughout many regions of the brain. Is all learning reducible to a single common form? Neuropsychological dissociations suggest that the mammalian brain possesses a number of different and potentially independent memory systems, with different mechanisms and anatomical dispositions, some of which are neurally widely dispersed and others of which are narrowly organized. Among the types considered are: (i) short-term memory; (ii) knowledge and skills; (iii) stable associative memory; (iv) event memory; and (v) priming. As double or multiple dissociations do not lead to logically inevitable conclusions, it has been argued that an alternative to multiple memory systems is variable modes of processing. But these, too, would be dissociable on the same lines of evidence. Dissociations, if strong and absolute, have strong pragmatic power when they are combined with evolutionary and neuroscientific evidence. Multiple memory systems may possibly share some common cellular mechanisms, but such mechanisms do not define the separate properties at the systems level.     

药物成瘾及成瘾记忆的研究现状

本文在介绍药物成瘾与学习和记忆密切相关的神经回路及共同分子机制的基础上,围绕学习和记忆在药物成瘾中的作用,综述了关联性学习与复吸,关联性学习与敏化,异常关联性学习与强迫性用药行为,关联性学习及成瘾记忆与成瘾,多重记忆系统与成瘾的发生发展等方面的研究进展,并强调了突触可塑性及成瘾记忆在药物成瘾中的重要性。在此基础上提出：作为慢性脑病的药物成瘾的形成过程的重要特征是它包含着信息的特殊学习类型。药物成瘾与依赖于多巴胺的关联性学习紊乱有密切关系。海马可能在成瘾中扮演重要角色。     

Spectral EEG characteristics in emotional stress model in cats

Emotional stress induced a global activation of the brain electrical activity: a decrease in the power of the theta-, alpha-, and beta-1-frequency bands in animals with active behaviour, and a decrease in the power of the theta- and alpha-frequency bands in animals with behavioural inhibition. The findings suggest that two forms of emotional behaviour correspond to different patterns of the brain activity. Appearance of spindle-like activity in passive animals' parietal and occipital leads suggests a hypothetical suppression in associative parietal and visual neocortical areas involved in processing the sensory information.     

Latent foci of excitation and the unpredictability of behavioral reactions

As a model for studying of "subconsciousness" mechanisms, latent foci of excitation may serve formed in animals under different influences on the brain: direct current, endogenous shifts and also under trace excitation after suprathreshold activity. It is experimentally shown that latent foci of excitation can intensify at activation of other CNS areas; as a result externally unmotivated "unpredictable" behavioural reaction arises. Between two latent foci of excitation associative connection can be established. The latent foci of excitation in the human CNS are also able to be activated under the influence of sensory stimuli, and behavioural reaction arising as a result of summation has no reflection in consciousness. Conclusion is made: it is rightful to consider the dominant foci of excitation formed at the level of subconsciousness as one of possible mechanisms of "unpredictability" of behaviour.     

Genetic control of processes of bacterial interactions with plants in associations

This review is devoted to the mechanisms and genetic control of processes underlying the formation and efficiency of associative relationships between bacteria and plants. The role of different polysaccharides and cellular fibrils in the appearance of associative relations and the biosynthetic pathway of these compounds and structures is considered. The molecular mechanisms of bacterial systems responsible for stimulating plant growth and development--nitrogen fixation and synthesis of a plant hormone, indoleacetic acid--are presented. The properties of associative bacteria are discussed in comparison with the relevant characteristics of the most studied free-living or symbiotic model species of bacteria.     

Cytokine neurobiology: behavior and adaptive responses

Reactions of the brain to systemic LPS or IL-1 beta treatment were shown to have different thresholds, be mediated by different neurotransmitter systems, and have different mechanisms of realisation. Changes in behaviour and neurotransmitter systems activity of the hypothalamus induced by a systemic IL-1 beta treatment were shown to be mediated by its receptors in the brain. Expression of mRNA of the tumour necrosis factor was revealed in the rabbit brain following administration of high pyrogenic doses of the LPS. The data obtained corroborate the concept of the cytokines role in maintenance of the defence responses in activation of the immune system, as well as their probable role in normal mechanisms of physiological functions control.     

Neuronal signalling of fear memory

The learning and remembering of fearful events depends on the integrity of the amygdala, but how are fear memories represented in the activity of amygdala neurons? Here, we review recent electrophysiological studies indicating that neurons in the lateral amygdala encode aversive memories during the acquisition and extinction of Pavlovian fear conditioning. Studies that combine unit recording with brain lesions and pharmacological inactivation provide evidence that the lateral amygdala is a crucial locus of fear memory. Extinction of fear memory reduces associative plasticity in the lateral amygdala and involves the hippocampus and prefrontal cortex. Understanding the signalling of aversive memory by amygdala neurons opens new avenues for research into the neural systems that support fear behaviour.     

Integrity principle of the evolution of associative centers in the vertebrate brain

Principle trends in the evolution of brain associative centers are discussed. It is demonstrated that in various taxonomic groups the possibilities of adaptive changes, as far as the brain is concerned, go beyond the scope of biological, topographical, and dynamic coordinations, typical for other scopes and systems. The reason for such peculiarity of brain evolution is the multifunctional nature of nerve tissue. This property of nerve tissue allows vertebrates to implement the integrity principle and still preserve the flexibility of the brain, which is highly specialized morphologically and functionally.     

Parasitoidism, not sociality, is associated with the evolution of elaborate mushroom bodies in the brains of hymenopteran insects

The social brain hypothesis posits that the cognitive demands of social behaviour have driven evolutionary expansions in brain size in some vertebrate lineages. In insects, higher brain centres called mushroom bodies are enlarged and morphologically elaborate (having doubled, invaginated and subcompartmentalized calyces that receive visual input) in social species such as the ants, bees and wasps of the aculeate Hymenoptera, suggesting that the social brain hypothesis may also apply to invertebrate animals. In a quantitative and qualitative survey of mushroom body morphology across the Hymenoptera, we demonstrate that large, elaborate mushroom bodies arose concurrent with the acquisition of a parasitoid mode of life at the base of the Euhymenopteran (Orussioidea + Apocrita) lineage, approximately 90 Myr before the evolution of sociality in the Aculeata. Thus, sociality could not have driven mushroom body elaboration in the Hymenoptera. Rather, we propose that the cognitive demands of host-finding behaviour in parasitoids, particularly the capacity for associative and spatial learning, drove the acquisition of this evolutionarily novel mushroom body architecture. These neurobehavioural modifications may have served as pre-adaptations for central place foraging, a spatial learning-intensive behaviour that is widespread across the Aculeata and may have contributed to the multiple acquisitions of sociality in this taxon.     

Neuroendocrine and neurochemical mechanisms of the domestication of animals

A review of experimental data documenting that domestication of animals is associated with hereditary reorganization of neuro-endocrine mechanisms, responsible for basic processes of ontogeny, is presented. The data demonstrated changes in gonadal and pituitary-adrenal systems in domesticated animals. Analysis of evidence that selection for low aggressiveness towards man is, in fact, the selection for definite activity of brain neurotransmitters regulating aggressive behaviour and emotional stress response has been carried out. Supposed role of modifications in the mechanisms of domestication is discussed.     

Further exploration into the adaptive design of the arthropod "microbrain": I. Sensory and memory-processing systems

Arthropods have small but sophisticated brains that have enabled them to adapt their behavior to a diverse range of environments. In this review, we first discuss some of general characteristics of the arthropod "microbrain" in comparison with the mammalian "megalobrain". Then we discuss about recent progress in the study of sensory and memory-processing systems of the arthropod "microbrain". Results of recent studies have shown that (1) insects have excellent capability for elemental and context-dependent forms of olfactory learning, (2) mushroom bodies, higher olfactory and associative centers of arthropods, have much more elaborated internal structures than previously thought, (3) many genes involved in the formation of basic brain structures are common among arthropods and vertebrates, suggesting that common ancestors of arthropods and vertebrates already had organized head ganglia, and (4) the basic organization of sensori-motor pathways of the insect brain has features common to that of the mammalian brain. These findings provide a starting point for the study of brain mechanisms of elaborated behaviors of arthropods, many of which remain unexplored.     

Molecular-chemical basis of emotional states and reinforcement

In this paper we have attempted to relate and explain the activity of the brain different integrative levels, which ends up with occurrence of a goal-directed behaviour and reinforcement. Two main mechanisms of responding to subjects and events of external environment, as rapid executive reactions and slow motivational-emotional states, have been marked out. Each mechanism gets its own neurochemical basis, ways of information transmission and molecular-genetic programs for the specialized metabolic reactions. Together both mechanisms form a unique spatial-temporal continuum of neural processes aimed to produce the beneficial activity and reinforce it by subjectively perceived emotional states.     

Genesis of different components of evoked response in the associative parietal cortex of the cat's brain

The responses evoked in the associative parietal region of the brain by different peripheral stimuli constitute a complex reaction. An early short-latent component of the associative response (ECAR) is formed as the result of specific afferentation, whereas the long-latent late component that follows it is evoked by pulsations from nonspecific, reticular, and associative nuclei in the thalamus or by pulsations arriving from specific nuclei by indirect routes. The cortical mechanisms whereby ECAR arise may be provided by the structural-organization characteristics of interneuronal connections, in particular by the presence in all cortical layers of the terminals of afferent fibers, which establish contact that are mainly axodendritic.Institute of Normal and Pathological Physiology, Academy of Medical Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 2, No. 4, pp. 399–405, July–August, 1970.     

Looking for sexual selection in the female brain

Female mate choice behaviour has significant evolutionary consequences, yet its mechanistic origins are not fully understood. Recent studies of female sensory systems have made great strides in identifying internal mechanisms governing female preferences. Only recently, however, have we begun to identify the dynamic genomic response associated with mate choice behaviour. Poeciliids provide a powerful comparative system to examine genomic responses governing mate choice and female preference behaviour, given the great range of mating systems: from female mate choice taxa with ornamental courting males to species lacking male ornamentation and exhibiting only male coercion. Furthermore, they exhibit laboratory-tractable preference responses without sexual contact that are decoupled from reproductive state, allowing investigators to isolate mechanisms in the brain without physiological confounds. Early investigations with poeciliid species (Xiphophorus nigrensis and Gambusia affinis) have identified putative candidate genes associated with female preference response and highlight a possible genomic pathway underlying female social interactions with males linked functionally with synaptic plasticity and learning processes. This network is positively correlated with female preference behaviour in the female mate choice species, but appears inhibited in the male coercive species. This behavioural genomics approach provides opportunity to elucidate the fundamental building blocks, and evolutionary dynamics, of sexual selection.     

Spatial inference without a cognitive map: the role of higher-order path integration

The cognitive map has been taken as the standard model for how agents infer the most efficient route to a goal location. Alternatively, path integration – maintaining a homing vector during navigation – constitutes a primitive and presumably less-flexible strategy than cognitive mapping because path integration relies primarily on vestibular stimuli and pace counting. The historical debate as to whether complex spatial navigation is ruled by associative learning or cognitive map mechanisms has been challenged by experimental difficulties in successfully neutralizing path integration. To our knowledge, there are only three studies that have succeeded in resolving this issue, all showing clear evidence of novel route taking, a behaviour outside the scope of traditional associative learning accounts. Nevertheless, there is no mechanistic explanation as to how animals perform novel route taking. We propose here a new model of spatial learning that combines path integration with higher-order associative learning, and demonstrate how it can account for novel route taking without a cognitive map, thus resolving this long-standing debate. We show how our higher-order path integration (HOPI) model can explain spatial inferences, such as novel detours and shortcuts. Our analysis suggests that a phylogenetically ancient, vector-based navigational strategy utilizing associative processes is powerful enough to support complex spatial inferences.