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The development of embodied cognition: six lessons from babies.

The embodiment hypothesis is the idea that intelligence emerges in the interaction of an agent with an environment and as a result of sensorimotor activity. We offer six lessons for developing embodied intelligent agents suggested by research in developmental psychology. We argue that starting as a baby grounded in a physical, social, and linguistic world is crucial to the development of the flexible and inventive intelligence that characterizes humankind.

Artificial Intelligence↗

Neurobiological effects of a null mutation depend on genetic context: comparison between two hotfoot alleles of the delta-2 ionotropic glutamate receptor.

Hotfoot is a mutant mouse with an ataxic phenotype which has been shown to be due to a mutation in the Grid2 gene. In this paper, we compare molecular, morphological, electrophysiological and behavioral features of two Grid2 alleles: Grid2(ho-4J) and Grid2(ho-Nancy). We first show that these two mutations are deletions in the open reading frame of the gene and that no GRID2 protein is detectable in extracts of mutant cerebella, suggesting that the two alleles are null-like mutations. Morphological and electrophysiological analyses reveal no obvious differences between the two strains: both strains showed the naked Purkinje dendritic spines and mismatch between the length of the presynaptic active zone and postsynaptic differentiation characteristic of the hotfoot mutation; and the same low level (20%) of multiple climbing fiber innervation of Purkinje cells was found in both strains. Only differences in motor behavior were found between the two strains. The Grid2(ho-4J) mouse shows more severe ataxia that the Grid2(ho-Nancy) mouse and, although both strains show a clear capacity to improve their performance of a motor task with training, the Grid2(ho-4J) performance remains very poor whereas Grid2(ho-Nancy) mice approach control levels. The only difference between the two strains is their genetic background. Our results show that the genetic background must be taken into account when analyzing sensorimotor performances of mutant mice.

Afferent Pathways↗

Striatal level of regulation of learned forepaw movements in rats.

The role of the striatal adenylyl cyclase (AC) and cholinergic systems in the learning and expression of new forepaw movements (reaching with prolonged pushing on a fixed piston) was studied in male Wistar rats. Motor learning processes, prenatal hypoxia, and cholinergic drugs changed the properties of the AC system in the striatum. After learning, the striatal basal AC activity was decreased compared to untrained control rats. In addition, the AC activity was more decreased in animals with a good ability to learn compared to poor learners (up to 31 % and 51 %, correspondingly; p<0.01). Rats subjected to prenatal hypoxia (13-14th days of embryogenesis) had a lower ability to learn the new movements requiring tactile control and the striatal AC activity in these rats was 1.8 times higher (p<0.001) than controls. In vitro application of the cholinergic agonist carbachol (CARB) 10-5 M (corresponding to approximately 0.3 microg), as well as the antagonist scopolomine (SCOP) 10(-5) M (approximately 0.3 microg) decreased AC activity in the synaptosomal fraction of the striatum. In vivo injections of CARB (0.3-3 microg/1microl) or SCOP (0.3-3 microg/1microl) into the ventral striatum (nucleus accumbens) modified the newly learned sensorimotor skill. After CARB injections the rats performed slower movements with more prolonged pushing. After SCOP the rats could not retain the learned pushing movement. These in vivo and in vitro data suggest that the cholinergic mediator system of the striatum is involved in learning sensory-controlled forepaw movements as well as the regulation of new motor skills by modulating the AC signal transduction process in the striatum. The data confirmed that modification of the striatal AC system resulted in the modulation of reaching behavior and better expression of the learned reflex.

Adenylyl Cyclase Inhibitors↗

Functional assessments in mice and rats after focal stroke.

This paper presents a comprehensive assessment of sensorimotor deficits in the mouse after focal ischaemia induced by occlusion of the middle cerebral artery. Twenty four hours after induction of middle cerebral artery occlusion, mice showed deficits in a range of sensory and motor tasks as assessed by the SHIRPA protocol. In addition they exhibited a decrease in rotarod performance and locomotor activity. Some behaviours, such as locomotor activity, were also impaired in sham operated animals compared to normal controls, although these impairments were not as marked as those exhibited by the ischaemic mice. This is the first comprehensive analysis of the short term effects of permanent focal ischaemia in mice. In a second series of experiments in the rat, rates of recovery over time were examined. Simple (neurological grades, rotarod) and complex (sticky label test) tasks were examined in rats after middle cerebral artery occlusion up to 7 days post-ischaemia. Ischaemic rats had a profound deficit in contralateral performance on the sticky label task with no evidence of recovery. A less marked deficit was also observed in ipsilateral performance of this task. These deficits were still present 7 days after ischaemia. Ischaemic rats also exhibited a deficit on rotarod performance but this had recovered 7 days post-ischaemia. Thus different sensorimotor tasks have different rates of recovery after focal cerebral ischaemia in the rat. Further characterisation of these tasks will enhance their utility meaningful preclinical means of assessing functional recovery of the administration of potential neuroprotective and regenerative therapies.

Animals↗

Brain correlates of fast and slow handwriting in humans: a PET-performance correlation analysis.

The present study examined the cerebral control of velocity during handwriting. We employed H215O positron emission tomography (PET) to measure the regional cerebral blood flow (rCBF) in 10 healthy subjects. Participants were required to write the German verb 'bellen' ('to bark') either at their normal speed (i.e. fast open-loop handwriting) or to write at approximately half of their normal speed without visual feedback. The second task required a continuous modification of the motor output according to the kinaesthetic feedback from the hand (i.e. slow closed-loop handwriting). Pencil movements were recorded during PET scanning and analysed off-line using a stroke-based analysing program. The mean number of inversions in velocity (NIV) per stroke was used to quantify the mode of motor control during each PET scan. A NIV of 1 indicates fast open-loop processing, whereas an increase in NIV reflects a shift towards slow closed-loop processing of handwriting. Foci in the left primary sensorimotor cortex, the right lateral premotor cortex, the left anterior parietal cortex, the left anterior putamen, the left rostral supplementary motor area and the right precuneus showed a graded increase in functional activation with the mean NIV per stroke, suggesting that this set of brain regions is particularly involved in the processing of slow closed-loop writing movements. No area showed a negative relationship between rCBF and the mean NIV per stroke, suggesting that fast open-loop handwriting is achieved by an optimized cooperation of the manual sensorimotor network rather than by a selective activation of a distinct network component.

Adult↗

[The functional bases for the formation of sensory system interaction in ontogeny].

The study of the mechanisms of the intersensory interaction and its development in the postnatal ontogenesis. The first proved evidence of participation of the intracortical inhibition mechanisms in the processes of heterosensory interaction on the cortical level. The study of formation in kittens of the heterosensory interaction on the neurons of the sensorimotor cortex area has established the correlation of the process with the interconnected maturing of the afferent entries to this cortex area, and the system of intracortical inhibition. Correlation was established between the time of development of the heterosensory interactions on neurons of the sensorimotor cortex area in ontogenesis and the age period when injury of the sensorimotor cortex destroys the ability of animals to produce a conditioned reflex to a complex heteromodal signal in case of inhibiting reactions to its components. Discussion of the problem of input of the cortical level to the process of heterosensory interaction in different phases of ontogenesis.

Aging↗

Synaptic reorganization in the hippocampal formation of alcohol-fed rats may compensate for functional deficits related to neuronal loss.

We have examined the behavioral and neuroanatomical effects of long-term alcohol intake in rats ingesting a 20% solution of ethanol for 30 weeks. Previous studies have shown that this treatment provokes neuronal degeneration in the hippocampal formation, which occurs in parallel with remodeling processes. Spatial reference and working memory of alcohol-fed rats were evaluated during last 4 weeks of treatment by comparison of their performance with age-matched controls on the Morris water maze. Alcohol consumption did not affect the performance of rats in the reference memory task as indicated by the measures derived from the acquisition trials and from the probe-trial, which were highly similar for alcohol-fed and control animals. Also, performance in the working memory task was not significantly altered in alcohol-treated animals. No treatment-related changes in swim speed or impairments of sensorimotor abilities, tested in the visible platform task, were detected. Stereological methods were applied to evaluate the damage inflicted by alcohol intake in the structure of the hippocampal formation. In the alcohol-treated animals, there was a noticeable cell loss in the granular layer of the dentate gyrus (10%), and in CA3 (18%) and CA1 (19%) hippocampal subdivisions. In spite of the neuronal loss, the total number of synapses between mossy fibers and CA3 pyramids was unaffected by alcohol treatment suggesting that new synaptic contacts were formed between the surviving neurons. We show that, regardless the marked hippocampal cell loss in rats exposed to chronic alcohol intake, the reorganization that takes place at the synaptic level may alleviate the expected functional deficits.

Alcohol Drinking↗

Cysteamine impairs the development of the acoustic startle response in rats: possible role of somatostatin.

Somatostatin is transiently expressed in many regions of the developing brain, among others in auditory brainstem nuclei of neonatal rats. To explore the functional significance of somatostatin during the ontogeny of acoustically elicited behavior, the acoustic startle response (ASR) was measured in developing rats after chronic application of cysteamine, which, when applied in low doses, most selectively depletes somatostatin. Cysteamine treatment drastically reduced somatostatin immunoreactivity in the cochlear nuclear complex and the caudal pontine reticular nucleus, i.e. in structures mediating the ASR. It did not affect the ASR amplitude of postnatal day (P) 13 animals, yet it resulted in a significant reduction of the ASR amplitude at P18. Our results therefore suggest that somatostatin can influence the maturation of sensorimotor information processing.

Acoustic Stimulation↗

Local as well as remote functional and metabolic changes after focal ischemia in cats.

Behavior and limb placing ability were analyzed acutely and subacutely (up to 21 days) following unilateral occlusion of the middle cerebral artery (MCA) in cats. Immediately following occlusion, all tested cats started to display a sequence of different behaviors, characteristic for 1) an ipsilateral inhibition of dopaminergic activity in the caudate nucleus (CN); 2) an inhibition of GABAergic activity in the reticular substantia nigra (SNR); 3) a stimulation of GABA receptors in the deeper layers of the colliculus superior (CSDL) (starting-time of these phases: about 4, 12 and 25 min, respectively). The latter behavior was also present subacutely. In addition, unilateral orofacial dyskinetic movements were observed acutely as well as subacutely. Contralateral limb placing was deficient in all cats 60 min postocclusion; it was at least partly restored subacutely. Twenty-one days after the occlusion, [14C]-2-D-deoxyglucose uptake was relatively reduced in the ipsilateral CN (especially in its posterior part), the ipsilateral SNR and the ipsilateral CSDL. The anterior CN appeared to be less affected than the posterior CN. Metabolism was relatively reduced in the sensorimotor cortex only in part of the tested cats. The data show that unilateral MCA occlusion produces consistent functional changes in all structures studied apart from the sensorimotor cortex, viz. the CN, the SNR and the CSDL.

Animals↗

Progressive degeneration of dopamine system functions after transient cerebral oligemia in rats.

A reduction in cerebral blood flow to oligemic levels was achieved in pentobarbital-anesthetized adult rats by clamping both carotid arteries (BCCA) for 60 min. To assess the extent to which the animals' dopaminergic system was affected over an increasing time span, their spontaneous locomotor activity in an unfamiliar environment and in response to the subcutaneous administration of apomorphine was tested at various times after either BCCA or sham operation. Eight to 14 days after the operation, it was possible to observe a diminished locomotor activity in response to apomorphine injection in BCCA as compared with sham-operated animals, while oral stereotypical behavior such as licking was increased. At 3 months, there was only a subtle decrease in apomorphine-induced locomotor activity, and stereotypical behavior was similar in both groups. At 7 months, the BCCA rats covered shorter distances than sham-operated controls during the habituation phase; after apomorphine injection, more stereotypic movements, such as, e.g., sniffing, were observed, and less running. Twelve months after surgery, no further differences could be observed between the two groups during the habituation phase, but the injection of apomorphine led to increased stereotypic sniffing movements, rearing and locomotor activity in BCCA animals to a greater extent than in the controls. At 12 months, sensorimotor disturbances elicited by the rota rod test, which were only transiently observed at 11 weeks and 7 months, did not appear any different from the normal age-related motor decline of the sham-operated controls. The animals' motor co-ordination in the chimney test was not significantly disturbed during the time between 7 and 12 months after surgery. At 15 months, nocturnal locomotor activities in BCCA rats were significantly decreased. In situ hybridization (ISH) histochemistry revealed decreased D1 receptor mRNA (D1RmRNA) in striatal neurons 19 months after surgery, while D2 receptor mRNA (D2RmRNA) and the neuronal number remained the same. The present results show that just as is already known for the immature rat brain, the adult rat brain, too, reacts to a transient decrease in its blood supply by appearance of long-lasting alterations in function, and that even a single oligemic episode is capable of inducing progressive dopaminergic dysfunctions and ultimately the partial loss of striatal D1RmRNA.

Animals↗

An analysis of feeding and sensorimotor abilities of rats after decortication.

Feeding and sensorimotor abilities of decorticate rats are described. Within 24 hr after surgery, the rats ate mash, unfamiliar foods, dry chow, and tap and sugar water from a spatula. They sampled nonnutritive substances but rejected noxious substances. Although initially anorectic, given mash, they arrested weight loss as early as 2 to 4 days. With training, they drank water from a spout by Day 10 and gained weight on dry food and water. Throughout survival they had difficulties with chewing food, lapping water, and trimming toenails. They showed sensorimotor neglect to objects touched to the body; they did not orient to the place in space where touched, they did not orient when touched on the head. There may be different types of orienting, one which is lost and one spared following decortication. When lifted from a surface, they showed limb and postural abnormalities; when dropped, righting responses were impaired. They showed stereotyped reflexive responses to surface and became trapped on platforms and in alleys. The range of abnormalities, from simple acts to more complex behaviors, considered within the context of feeding behavior in its broadest sense, shows the rats simplest environment. Furthermore, cortically controlled feeding seems partly independent from hypothalamically controlled feeding.

Animals↗

Review of the NIMH Israeli Kibbutz-City Study and the Jerusalem Infant Development Study.

The National Institute of Mental Health (NIMH) Israeli Kibbutz-City Study has followed the development of offspring of schizophrenic parents from middle childhood through early adulthood. During childhood, a subgroup of offspring of schizophrenic patients showed clear neurobehavioral deficits often accompanied by poor social competence. Early followup data suggest that this subgroup of high-risk children is at greatest risk for adult schizophrenia spectrum illness. The Jerusalem Infant Development Study has followed a similar population of children at risk for schizophrenia from before birth through middle childhood. A subgroup of dysfunctioning in the high-risk children showed sensorimotor dysfunctioning in the first year of life, which was followed by perceptual, motor, and attentional dysfunctioning in childhood--identical to that found in the NIMH cohort. Results from both studies support the hypothesis that schizophrenic illness involves constitutional factors whose expression can be observed as early as infancy. Results also illustrate the importance of using data-analytic approaches that (1) look for subgroups within high-risk groups rather than only group differences between high- and low-risk groups, and (2) examine profiles of behavior rather than only single variables.

Adolescent↗

Electropalatographic, acoustic, and perceptual data on adaptation to a palatal perturbation.

Exploring the compensatory responses of the speech production system to perturbation has provided valuable insights into speech motor control. The present experiment was conducted to examine compensation for one such perturbation-a palatal perturbation in the production of the fricative /s/. Subjects wore a specially designed electropalatographic (EPG) appliance with a buildup of acrylic over the alveolar ridge as well as a normal EPG palate. In this way, compensatory tongue positioning could be assessed during a period of target specific and intense practice and compared to nonperturbed conditions. Electropalatographic, acoustic, and perceptual analyses of productions of /asa/ elicited from nine speakers over the course of a one-hour practice period were conducted. Acoustic and perceptual results confirmed earlier findings, which showed improvement in production with a thick artificial palate in place over the practice period; the EPG data showed overall increased maximum contact as well as increased medial and posterior contact for speakers with the thick palate in place, but little change over time. Negative aftereffects were observed in the productions with the thin palate, indicating recalibration of sensorimotor processes in the face of the oral-articulatory perturbation. Findings are discussed with regard to the nature of adaptive articulatory skills.

Adult↗

[Psychological measurement of tactile-kinesthetic perception in early childhood].

The perception theory of Affolter (1987), the theory of sensory integration of Ayres (1979, 1984) as well as the development theory of Piaget (1973) consider sensorimotor experiences as a basis for the child's cognitive development. Tactile-kinesthetic perception has hereby a central position. In the German-speaking psychology, no standardized tests exist to measure the developmental age of tactile-kinesthetic perception in early childhood. The subtests of neuropsychological batteries have not been primarily constructed for young children, therefore they may not portray the age dependence of tactile-kinesthetic perception exactly. That is why we have collected a pool of items, empirically based descriptors of tactile-kinesthetic behavior, to test this perceptual modality. Then we proved it in a series of pretests. The resulting preliminary developmental test contains 7 functions: Localization of touch, pressure sensibility, two-point-discrimination, finger identification, object stereognosis, stereognosis of object properties, and graphesthesia. It was administered to 111 children aged from 3;2 to 6;5 years in kindergarten. Data of the item analysis demonstrated that many items proved to be to easy. Nevertheless, the instrument is useful and measures reliably. The deficits of the experimental test version will now be corrected by a revision. We will start a detailed analysis again using the revised test.

Child↗

IL-2/15 receptor-beta gene deletion alters neurobehavioral performance.

The common IL-2/15 receptor-beta (IL-2/15Rbeta) is an essential signaling subunit that is shared exclusively by IL-2 and IL-15, and is enriched in the hippocampal formation and related limbic regions. We have previously shown that mice lacking IL-2 exhibit alterations in hippocampal-dependent learning, sensorimotor gating and accompanying reductions in hippocampal infrapyramidal mossy neuronal fiber length. Although the effects of exogenous IL-2 on various aspects of forebrain neuronal function are well documented, it is unclear whether IL-15 has neuromodulatory actions. Here we sought to test the hypothesis that the combined loss of the ability of IL-2 and IL-15 to signal through IL-2/15Rbeta in the brain would influence neurobehavioral performance, in particular spatial learning and memory performance. To test this hypothesis, we compared several different domains of behavior in mice that had one or both IL-2/15Rbeta gene alleles deleted. Compared with C57BL/6-IL-2/15Rbeta+/+ wild-type and C57BL/6-IL-2/15Rbeta+/- heterozygote littermates, C57BL/6-IL-2/15Rbeta-/- knockout mice exhibited a deficit in prepulse inhibition of the acoustic startle reflex (PPI). The IL-2/15Rbeta knockout mice also showed significant reductions in acoustic startle reactivity, and modest differences in behavior in the elevated plus-maze test indicative of reduced levels of fearfulness in response to novelty. The IL-2/15Rbeta knockout mice did not differ in locomotor activity in either the plus-maze or the Morris water-maze, and contrary to our working hypothesis, they did not differ in spatial learning or memory performance in the water-maze. Further studies are required to determine if these behavioral alterations may be attributable to factors such as the loss of the ability of IL-15 and/or IL-2 to modulate limbic neurons, autoimmunity or genetic factors associated with IL-2/15Rbeta gene deletion.

Acoustic Stimulation↗

Three-dimensional eye-head coordination is implemented downstream from the superior colliculus.

How the brain transforms two-dimensional visual signals into multi-dimensional motor commands, and subsequently how it constrains the redundant degrees of freedom, are fundamental problems in sensorimotor control. During fixations between gaze shifts, the redundant torsional degree of freedom is determined by various neural constraints. For example, the eye- and head-in-space are constrained by Donders' law, whereas the eye-in-head obeys Listing's law. However, where and how the brain implements these laws is not yet known. In this study, we show that eye and head movements, elicited by unilateral microstimulations of the superior colliculus (SC) in head-free monkeys, obey the same Donders' strategies observed in normal behavior (i.e., Listing's law for final eye positions and the Fick strategy for the head). Moreover, these evoked movements showed a pattern of three-dimensional eye-head coordination, consistent with normal behavior, where the eye is driven purposely out of Listing's plane during the saccade portion of the gaze shift in opposition to a subsequent torsional vestibuloocular reflex slow phase, such that the final net torsion at the end of each head-free gaze shift is zero. The required amount of saccade-related torsion was highly variable, depending on the initial position of the eye and head prior to a gaze shift and the size of the gaze shift, pointing to a neural basis of torsional control. Because these variable, context-appropriate torsional saccades were correctly elicited by fixed SC commands during head-free stimulations, this shows that the SC only encodes the horizontal and vertical components of gaze, leaving the complexity of torsional organization to downstream control systems. Thus we conclude that Listing's and Donders' laws of the eyes and head, and their three-dimensional coordination mechanisms, must be implemented after the SC.

Algorithms↗

Region-specific sprouting of crossed corticofugal fibers after unilateral cortical lesions in adult mice.

Long considered to be limited to early development or restricted adult brain regions in mammals, axonal sprouting of spared axons into denervated brain areas now appears more widespread in the adult mammalian brain. However, its extent and mechanisms remain poorly understood. In this study, we show that robust sprouting of corticofugal axons occurs in the dorsolateral striatum but not the red nucleus of adult mice after unilateral lesions of the sensorimotor cortex induced either by mechanical removal or by thermocoagulation of pial blood vessels. These results show that local factors are critical for axonal sprouting in adult brain. They also extend previous findings in rats to a species readily amenable to genetic analysis in order to elucidate the mechanisms of this effect.

Animals↗

On feeling and being felt with.

The development of feelings in early childhood is traced, with special emphasis on the mother's role during the toddler phase when she facilitates her child's transition from sensorimotoric discharge to the mental experience, ownership, and use of modulated affects. Many child analytic patients use defenses to ward off feelings, many have not even reached the developmental level of experiencing feelings. How this difficulty manifests itself, the reasons for the developmental lag, and the analytic means of helping such patients are discussed and illustrated.

Child Behavior Disorders↗