Cortical asymmetries at different spatial hierarchies relate to phonological processing ability

The ability to map speech sounds to corresponding letters is critical for establishing proficient reading. People vary in this phonological processing ability, which has been hypothesized to result from variation in hemispheric asymmetries within brain regions that support language. A cerebral lateralization hypothesis predicts that more asymmetric brain structures facilitate the development of foundational reading skills like phonological processing. That is, structural asymmetries are predicted to linearly increase with ability. In contrast, a canalization hypothesis predicts that asymmetries constrain behavioral performance within a normal range. That is, structural asymmetries are predicted to quadratically relate to phonological processing, with average phonological processing occurring in people with the most asymmetric structures. These predictions were examined in relatively large samples of children (N = 424) and adults (N = 300), using a topological asymmetry analysis of T1-weighted brain images and a decoding measure of phonological processing. There was limited evidence of structural asymmetry and phonological decoding associations in classic language-related brain regions. However, and in modest support of the cerebral lateralization hypothesis, small to medium effect sizes were observed where phonological decoding accuracy increased with the magnitude of the largest structural asymmetry across left hemisphere cortical regions, but not right hemisphere cortical regions, for both the adult and pediatric samples. In support of the canalization hypothesis, small to medium effect sizes were observed where phonological decoding in the normal range was associated with increased asymmetries in specific cortical regions for both the adult and pediatric samples, which included performance monitoring and motor planning brain regions that contribute to oral and written language functions. Thus, the relevance of each hypothesis to phonological decoding may depend on the scale of brain organization.     

High-resolution fMRI maps of cortical activation in nonhuman primates: correlation with intrinsic signal optical images

One of the most widely used functional brain mapping tools is blood oxygen level-dependent (BOLD) functional magnetic resonance imaging (fMRI). This method has contributed to new understandings of the functional roles of different areas in the human brain. However, its ability to map cerebral cortex at high spatial (submillimeter) resolution is still unknown. Other methods such as single- and multiunit electrophysiology and intrinsic signal optical imaging have revealed submillimeter resolution of sensory topography and cortical columnar activations. However, they are limited either by spatial scale (electrophysiology characterizes only local groups of neurons) or by the inability to monitor deep structures in the brain (i.e., cortical regions buried in sulci or subcortical structures). A method that could monitor all regions of the brain at high spatial resolution would be ideal. This capacity would open the doors to investigating, for example, how networks of cerebral cortical columns relate to or produce behavior. In this article we demonstrate that, without benefit of contrast agents, at a magnetic field strength of 9.4 tesla, BOLD fMRI can reveal millimeter-sized topographic maps of digit representation in the somatosensory cortex of the anesthetized squirrel monkey. Furthermore, by mapping the "funneling illusion," it is possible to detect even submillimeter shifts in activation in the cortex. Our data suggest that at high magnetic field strength, the positive BOLD signal can be used to reveal high spatial resolution maps of brain activity, a finding that weakens previous notions about the ultimate spatial specificity of the positive BOLD signal.     

Resolving the dynamics of EEG generators by multichannel recordings

The voltage recorded over the cortex (ECoG) or over the scalp (EEG) is generated by currents derived from many sources called “generators”. Different patterns and amplitudes are observed in aroused, sleepy, epileptic or other brain states. Differences in amplitude are generally attributed to differences in synchrony among generators. The degree of EEG synchrony is measured by the correlation between electrodes placed over different cortical regions. We present a new way to quantitatively assess the degree of synchronization of these generators via multichannel recordings. We illustrate how situations where there are several groups of generators with different inter-group and intra-group synchronies can be analyzed. Finally, we present a way to identify the organization of groups exhibiting topographic organization. Although the model presented here is highly simplified, several methods are based on averaging activity over increasingly larger areas. These types of measurements may be applied as well to EEG and ECoG recordings.     

Auditory cortex mapmaking: principles, projections, and plasticity

Maps of sensory receptor epithelia and computed features of the sensory environment are common elements of auditory, visual, and somatic sensory representations from the periphery to the cerebral cortex. Maps enhance the understanding of normal neural organization and its modification by pathology and experience. They underlie the derivation of the computational principles that govern sensory processing and the generation of perception. Despite their intuitive explanatory power, the functions of and rules for organizing maps and their plasticity are not well understood. Some puzzles of auditory cortical map organization are that few complete receptor maps are available and that even fewer computational maps are known beyond primary cortical areas. Neuroanatomical evidence suggests equally organized connectional patterns throughout the cortical hierarchy that might underlie map stability. Here, we consider the implications of auditory cortical map organization and its plasticity and evaluate the complementary role of maps in representation and computation from an auditory perspective.     

Learning through maps: functional significance of topographic organization in primary sensory cortex.

The presence of "maps" in sensory cortex is a hallmark of the mammalian nervous system, but the functional significance of topographic organization has been called into question by physiological studies claiming that patterns of neural behavioral activity transcend topographic boundaries. This paper discusses recent behavioral and physiological studies suggesting that, when animals or human subjects learn perceptual tasks, the neural modifications associated with the learning are distributed according to the spatial arrangement of the primary sensory cortical map. Topographical cortical representations of sensory events, therefore, appear to constitute a true structural framework for information processing and plasticity.     

Asymmetry pays: visual lateralization improves discrimination success in pigeons

Functional cerebral asymmetries, once thought to be exclusively human, are now accepted to be a widespread principle of brain organization in vertebrates [1]. The prevalence of lateralization makes it likely that it has some major advantage. Until now, however, conclusive evidence has been lacking. To analyze the relation between the extent of cerebral asymmetry and the degree of performance in visual foraging, we studied grain-grit discrimination success in pigeons, a species with a left hemisphere dominance for visual object processing [2,3]. The birds performed the task under left-eye, right-eye or binocular seeing conditions. In most animals, right-eye seeing was superior to left-eye seeing performance, and binocular performance was higher than each monocular level. The absolute difference between left- and right-eye levels was defined as a measure for the degree of visual asymmetry. Animals with higher asymmetries were more successful in discriminating grain from grit under binocular conditions. This shows that an increase in visual asymmetry enhances success in visually guided foraging. Possibly, asymmetries of the pigeon's visual system increase the computational speed of object recognition processes by concentrating them into one hemisphere while preventing the other side of the brain from initiating conflicting search sequences of its own.     

The Characteristic and Changes of the Event-Related Potentials (ERP) and Brain Topographic Maps before and after Treatment with rTMS in Subjective Tinnitus Patients

Objectives

To compare the event-related potentials (ERPs) and brain topographic maps characteristic and change in normal controls and subjective tinnitus patients before and after repetitive transcranial magnetic stimulation (rTMS) treatment.

Methods and Participants

The ERPs and brain topographic maps elicited by target stimulus were compared before and after 1-week treatment with rTMS in 20 subjective tinnitus patients and 16 healthy controls.

Results

Before rTMS, target stimulus elicited a larger N1 component than the standard stimuli (repeating sounds)in control group but not in tinnitus patients. Instead, the tinnitus group pre-treatment exhibited larger amplitude of N1 in response to standard stimuli than to deviant stimuli. Furthermore tinnitus patients had smaller mismatch negativity (MMN) and late discriminative negativity (LDN)component at Fz compared with the control group. After rTMS treatment, tinnitus patients showed increased N1 response to deviant stimuli and larger MMN and LDN compared with pre-treatment. The topographic maps for the tinnitus group before rTMS -treatment demonstrated global asymmetry between the left and right cerebral hemispheres with more negative activities in left side and more positive activities in right side. In contrast, the brain topographic maps for patients after rTMS-treatment and controls seem roughly symmetrical. The ERP amplitudes and brain topographic maps in post-treatment patient group showed no significant difference with those in controls.

Conclusions

The characterical changes in ERP and brain topographic maps in tinnitus patients maybe related with the electrophysiological mechanism of tinnitus induction and development. It can be used as an objective biomarker for the evaluation of auditory central in subjective tinnitus patients. These findings support the notion that rTMS treatment in tinnitus patients may exert a beneficial effect.     

Investigations into the organization of information in sensory cortex

One might take the exploration of sensory cortex in the first decades of the last century as the opening chapter of modern neuroscience. The combined approaches of (i) measuring effects of restricted ablation on functional capacities, both in the clinic and the laboratory, together with (ii) anatomical investigations of cortical lamination, arealization, and connectivity, and (iii) the early physiological probing of sensory representations, led to a fundamental body of knowledge that remains relevant to this day. In our time, there can be little doubt that its organization as a mosaic of columnar modules is the pervasive functional property of mammalian sensory cortex [Brain 120 (1997) 701]. If one accepts the assertion that columns and maps must improve the functioning of the brain (why else would they be the very hallmark of neocortex?), then the inevitable question is: exactly what advantages do they permit? In this review of our recent presentation at the workshop on Homeostasis, plasticity and learning at the Institut Henri Poincaré, we will outline a systematic approach to investigating the role of modular, map-like cortical organization in the processing of sensory information. We survey current evidence concerning the functional significance of cortical maps and modules, arguing that sensory cortex is involved not solely in the online processing of afferent data, but also in the storage and retrieval of information. We also show that the topographic framework of primary sensory cortex renders the encoding of sensory information efficient, fast and reliable.     

Acting unilaterally: Why do animals with strongly lateralized brains behave differently than those with weakly lateralized brains?

Cerebral lateralization was once thought to be unique to humans, but is now known to be widespread among the vertebrates. Lateralization appears to confer cognitive advantages upon those that possess it. Despite the taxonomic ubiquity and described advantages of lateralization, substantial individual variation exists in all species. Individual variation in cerebral lateralization may be tied to individual variation in behaviour and the selective forces that act to maintain variation in behaviour may also act to maintain variation in lateralization. The mechanisms linking individual variation in the strength of cerebral lateralization to individual variation in behaviour remain obscure. We propose here a general hypothesis which may help to explain this link. We suggest that individuals with strong and weak lateralizations behave differently because of differences in the ability of one hemisphere to inhibit the functions of the other in each type of brain organization. We also suggest a specific mechanism involving the asymmetric epithalamic nucleus, the habenula. We conclude by discussing some predictions and potential tests of our hypothesis.     

Limbic frontal cortex in hominoids: A comparative study of area 13

The limbic frontal cortex forms part of the neural substrate responsible for emotional reactions to social stimuli. Area 13 is one of the cortical areas long known to be part of the posterior orbitofrontal cortex in several monkey species, such as the macaque. Its presence nevertheless in the human brain has been unclear, and the cortex of the frontal lobe of the great and lesser apes remains largely unknown. In this study area 13 was identified in human, chimpanzee, bonobo, gorilla, orangutan, and gibbon brains, and cortical maps were generated on the basis of its cytoarchitecture. Imaging techniques were used to characterize and quantify the microstructural organization of the area, and stereological tools were applied for estimates of the volume of area 13 in all species. Area 13 is conservative in its structure, and features such as size of cortical layers, density of neurons, and space available for connections are similar across hominoids with only subtle differences present. In contrast to the homogeneity found in its organization, variation is present in the relative size of this cortical area (as a percentage of total brain volume). The human and the bonobo include a complex orbitofrontal cortex and a relatively smaller area 13. On the contrary the orangutan stands out by having a shorter orbitofrontal region and a more expanded area 13. Differences in the organization and size of individual cortical areas involved in emotional reactions and social behavior can be related to behavioral specializations of each hominoid and to the evolution of emotions in hominids. Am J Phys Anthropol 106:129–155, 1998. © 1998 Wiley-Liss, Inc.     

Representational plasticity in the mammalian brain cortex. (Review article)

The epithelial receptors are represented in the mammalian brain cortex in a genetically defined, strictly regulated manner. Until the 1970s, the cortical maps and the wiring of the central nervous system were thought to be rather static and unchangeable. Subsequently, however, studies of sensory and motor cortical maps in particular genetic strains of animals and in animals with different perinatal or adult histories have revealed that the map organization can be modified at any time between conception and death. Especially studies of the effects of peripheral and central lesions and of perceptual learning on the sensory and motor cortical representations have had a dramatic effect in alerting neuroscientists and therapists to the reorganizational capacity of the adult brain. From a theoretical aspect, these changes in the cortical maps provide useful models for an understanding of the changes that can occur in the integrative functions of complex brain networks throughout life.     

The effect of intracortical competition on the formation of topographic maps in models of Hebbian learning

Correlation-based learning (CBL) models and self-organizing maps (SOM) are two classes of Hebbian models that have both been proposed to explain the activity-driven formation of cortical maps. Both models differ significantly in the way lateral cortical interactions are treated, leading to different predictions for the formation of receptive fields. The linear CBL models predict that receptive field profiles are determined by the average values and the spatial correlations of the second order of the afferent activity patterns, wheras SOM models map stimulus features. Here, we investigate a class of models which are characterized by a variable degree of lateral competition and which have the CBL and SOM models as limit cases. We show that there exists a critical value for intracortical competition below which the model exhibits CBL properties and above which feature mapping sets in. The class of models is then analyzed with respect to the formation of topographic maps between two layers of neurons. For Gaussian input stimuli we find that localized receptive fields and topographic maps emerge above the critical value for intracortical competition, and we calculate this value as a function of the size of the input stimuli and the range of the lateral interaction function. Additionally, we show that the learning rule can be derived via the optimization of a global cost function in a framework of probabilistic output neurons which represent a set of input stimuli by a sparse code. Received: 23 June 1999 / Accepted in revised form: 05 November 1999     

Lateralized behaviour of a non-social cichlid fish (Amatitlania nigrofasciata) in a social and a non-social environment

Cerebral lateralization, the partitioning of cognitive function preferentially into one hemisphere of the brain, is a trait ubiquitous among vertebrates. Some species exhibit population level lateralization, where the pattern of cerebral lateralization is the same for most members of that species; however, other species show only individual level lateralization, where each member of the species has a unique pattern of lateralized brain function. The pattern of cerebral lateralization within a population and an individual has been shown to differ based on the stimulus being processed. It has been hypothesized that sociality within a species, such as shoaling behaviour in fish, may have led to the development and persistence of population level lateralization. Here we assessed cerebral lateralization in convict cichlids (Amatitlania nigrofasciata), a species that does not shoal as adults but that shoals briefly as juveniles. We show that both male and female convict cichlids display population level lateralization when in a solitary environment but only females show population level lateralization when in a perceived social environment. We also show that the pattern of lateralization differs between these two tasks and that strength of lateralization in one task is not predictive of strength of lateralization in the other task.     

Functional organization of adult motor cortex is dependent upon continued protein synthesis

The functional organization of adult cerebral cortex is characterized by the presence of highly ordered sensory and motor maps. Despite their archetypical organization, the maps maintain the capacity to rapidly reorganize, suggesting that the neural circuitry underlying cortical representations is inherently plastic. Here we show that the circuitry supporting motor maps is dependent upon continued protein synthesis. Injections of two different protein synthesis inhibitors into adult rat forelimb motor cortex caused an immediate and enduring loss of movement representations. The disappearance of the motor map was accompanied by a significant reduction in synapse number, synapse size, and cortical field potentials and caused skilled forelimb movement impairments. Further, motor skill training led to a reappearance of movement representations. We propose that the circuitry of adult motor cortex is perpetually labile and requires continued protein synthesis in order to maintain its functional organization.     

Interpreting fMRI data: maps, modules and dimensions

Neuroimaging research over the past decade has revealed a detailed picture of the functional organization of the human brain. Here we focus on two fundamental questions that are raised by the detailed mapping of sensory and cognitive functions and illustrate these questions with findings from the object-vision pathway. First, are functionally specific regions that are located close together best understood as distinct cortical modules or as parts of a larger-scale cortical map? Second, what functional properties define each cortical map or module? We propose a model in which overlapping continuous maps of simple features give rise to discrete modules that are selective for complex stimuli.     

The emergence of somatotopic maps of the body in S1 in rats: the correspondence between functional and anatomical organization

Most of what we know about cortical map development and plasticity comes from studies in mice and rats, and for the somatosensory cortex, almost exclusively from the whisker-dominated posteromedial barrel fields. Whiskers are the main effector organs of mice and rats, and their representation in cortex and subcortical pathways is a highly derived feature of murine rodents. This specialized anatomical organization may therefore not be representative of somatosensory cortex in general, especially for species that utilize other body parts as their main effector organs, like the hands of primates. For these reasons, we examined the emergence of whole body maps in developing rats using electrophysiological recording techniques. In P5, P10, P15, P20 and adult rats, multiple recordings were made in the medial portion of S1 in each animal. Subsequently, these functional maps were related to anatomical parcellations of S1 based on a variety of histological stains. We found that at early postnatal ages (P5) medial S1 was composed almost exclusively of the representation of the vibrissae. At P10, other body part representations including the hindlimb and forelimb were present, although these were not topographically organized. By P15, a clear topographic organization began to emerge coincident with a reduction in receptive field size. By P20, body maps were adult-like. This study is the first to describe how topography of the body develops in S1 in any mammal. It indicates that anatomical parcellations and functional maps are initially incongruent but become tightly coupled by P15. Finally, because anatomical and functional specificity of developing barrel cortex appears much earlier in postnatal life than the rest of the body, the entire primary somatosensory cortex should be considered when studying general topographic map formation in development.     

Organizational levels of the cerebral cortex: An integrated model

We propose a theoretical model of the cerebral cortex which is based on its cellular components and integrates its different levels of organization: (1) cells have general adaptive and memorization properties; (2) cortical columns are repetitive interneuronal circuits which determine an adaptive processing specific to the cerebral cortex; (3) cortical maps effect selective combinations which are very efficient to learn basic behaviourial adaptations such as invariant recognition of forms, visually-guided hand movements, or execution of structured motor programs; (4) the network between cortical areas has a global architecture which integrates successive learning experiences into coherent functions such as the human language.     

Functional Cerebral Asymmetry and Principles of Analysis of Lexical and Grammatical Material

This article deals with the role of functional cerebral asymmetry in the analysis of lexical and grammatical material. The author examined healthy test subjects with various types of cerebral organization of speech activity; i.e., with speech lateralization in the left and right hemispheres and bilateral speech lateralization. The test subjects with speech lateralization in the left hemisphere and bilateral lateralization are shown to prefer the formal-grammatical principle of the classification of both lexemes (distinguishing antonyms and synonyms) and grammatical constructions (distinguishing active and passive sentences). The test subjects with speech lateralization in the right hemisphere fail to follow the formal-grammatical principle of the classification of either lexemes (synonyms disappear) or grammatical constructions (active and passive sentences are united). The data obtained show that the linguistic abilities are determined by the functional specialization of the hemispheres.     

Cerebral Asymmetry of fMRI-BOLD Responses to Visual Stimulation

Hemispheric asymmetry of a wide range of functions is a hallmark of the human brain. The visual system has traditionally been thought of as symmetrically distributed in the brain, but a growing body of evidence has challenged this view. Some highly specific visual tasks have been shown to depend on hemispheric specialization. However, the possible lateralization of cerebral responses to a simple checkerboard visual stimulation has not been a focus of previous studies. To investigate this, we performed two sessions of blood-oxygenation level dependent (BOLD) functional magnetic resonance imaging (fMRI) in 54 healthy subjects during stimulation with a black and white checkerboard visual stimulus. While carefully excluding possible non-physiological causes of left-to-right bias, we compared the activation of the left and the right cerebral hemispheres and related this to grey matter volume, handedness, age, gender, ocular dominance, interocular difference in visual acuity, as well as line-bisection performance. We found a general lateralization of cerebral activation towards the right hemisphere of early visual cortical areas and areas of higher-level visual processing, involved in visuospatial attention, especially in top-down (i.e., goal-oriented) attentional processing. This right hemisphere lateralization was partly, but not completely, explained by an increased grey matter volume in the right hemisphere of the early visual areas. Difference in activation of the superior parietal lobule was correlated with subject age, suggesting a shift towards the left hemisphere with increasing age. Our findings suggest a right-hemispheric dominance of these areas, which could lend support to the generally observed leftward visual attentional bias and to the left hemifield advantage for some visual perception tasks.