Homeostatic Feelings and the Emergence of Consciousness.

In this article, we summarize our views on the problem of consciousness and outline the current version of a novel hypothesis for how conscious minds can be generated in mammalian organisms. We propose that a mind can be considered conscious when three processes are in place: the first is a continuous generation of interoceptive feelings, which results in experiencing of the organism's internal operations; the second is the equally continuous production of images, generated according to the organism's sensory perspective relative to its surround; the third combines feeling/experience and perspective resulting in a process of subjectivity relative to the image contents. We also propose a biological basis for these three components: the peripheral and central physiology of interoception and exteroception help explain the implementation of the first two components, whereas the third depends on central nervous system integration, at multiple levels, from spinal cord, brainstem, and diencephalic nuclei, to selected regions of the mesial cerebral cortices.

Feelings Are the Source of Consciousness.

In this view, we address the problem of consciousness, and although we focus on its human presentation, we note that the phenomenon is present in numerous nonhuman species and use findings from a variety of animal studies to explain our hypothesis for how consciousness is made. Consciousness occurs when mind contents, such as perceptions and thoughts, are spontaneously identified as belonging to a specific organism/owner. Conscious minds are said to have a self that experiences mental events. We hypothesize that the automatic identification that associates minds and organisms is provided by a continuous flow of homeostatic feelings. Those feelings arise from the uninterrupted process of life regulation and correspond to both salient physiological fluctuations such as hunger, pain, well-being, or malaise, as well as to states closer to metabolic equilibrium and best described as feelings of life/existence, such as breathing or body temperature. We also hypothesize that homeostatic feelings were the inaugural phenomena of consciousness in biological evolution and venture that they were selected because the information they provided regarding the current state of life regulation conferred extraordinary advantages to the organisms so endowed. The "knowledge" carried by conscious homeostatic feelings provided "overt" guidance for life regulation, an advance over the covert regulation present in nonconscious organisms. Finally, we outline a mechanism for the generation of feelings based on a two-way interaction between interoceptive components of the nervous system and a particular set of nonneural components of the organism's interior, namely, viscera and circulating chemical molecules involved in their operations. Feelings emerge from this interaction as continuous and hybrid phenomena, related simultaneously to two series of events. The first is best described by the terms neural/representational/and mental and the second by the terms nonneural/visceral/and chemical. We note that this account offers a solution for the mind-body problem: homeostatic feelings constitute the "mental" version of bodily processes.

Childhood Music Training Induces Change in Micro and Macroscopic Brain Structure: Results from a Longitudinal Study.

Several studies comparing adult musicians and nonmusicians have shown that music training is associated with structural brain differences. It is not been established, however, whether such differences result from pre-existing biological traits, lengthy musical training, or an interaction of the two factors, or if comparable changes can be found in children undergoing music training. As part of an ongoing longitudinal study, we investigated the effects of music training on the developmental trajectory of children's brain structure, over two years, beginning at age 6. We compared these children with children of the same socio-economic background but either involved in sports training or not involved in any systematic after school training. We established at the onset that there were no pre-existing structural differences among the groups. Two years later we observed that children in the music group showed (1) a different rate of cortical thickness maturation between the right and left posterior superior temporal gyrus, and (2) higher fractional anisotropy in the corpus callosum, specifically in the crossing pathways connecting superior frontal, sensory, and motor segments. We conclude that music training induces macro and microstructural brain changes in school-age children, and that those changes are not attributable to pre-existing biological traits.

The Neurobiology of Agrammatic Sentence Comprehension: A Lesion Study.

Broca's area has long been implicated in sentence comprehension. Damage to this region is thought to be the central source of "agrammatic comprehension" in which performance is substantially worse (and near chance) on sentences with noncanonical word orders compared with canonical word order sentences (in English). This claim is supported by functional neuroimaging studies demonstrating greater activation in Broca's area for noncanonical versus canonical sentences. However, functional neuroimaging studies also have frequently implicated the anterior temporal lobe (ATL) in sentence processing more broadly, and recent lesion-symptom mapping studies have implicated the ATL and mid temporal regions in agrammatic comprehension. This study investigates these seemingly conflicting findings in 66 left-hemisphere patients with chronic focal cerebral damage. Patients completed two sentence comprehension measures, sentence-picture matching and plausibility judgments. Patients with damage including Broca's area (but excluding the temporal lobe; n = 11) on average did not exhibit the expected agrammatic comprehension pattern-for example, their performance was >80% on noncanonical sentences in the sentence-picture matching task. Patients with ATL damage ( n = 18) also did not exhibit an agrammatic comprehension pattern. Across our entire patient sample, the lesions of patients with agrammatic comprehension patterns in either task had maximal overlap in posterior superior temporal and inferior parietal regions. Using voxel-based lesion-symptom mapping, we find that lower performances on canonical and noncanonical sentences in each task are both associated with damage to a large left superior temporal-inferior parietal network including portions of the ATL, but not Broca's area. Notably, however, response bias in plausibility judgments was significantly associated with damage to inferior frontal cortex, including gray and white matter in Broca's area, suggesting that the contribution of Broca's area to sentence comprehension may be related to task-related cognitive demands.

Processing Narratives Concerning Protected Values: A Cross-Cultural Investigation of Neural Correlates.

Narratives are an important component of culture and play a central role in transmitting social values. Little is known, however, about how the brain of a listener/reader processes narratives. A receiver's response to narration is influenced by the narrator's framing and appeal to values. Narratives that appeal to "protected values," including core personal, national, or religious values, may be particularly effective at influencing receivers. Protected values resist compromise and are tied with identity, affective value, moral decision-making, and other aspects of social cognition. Here, we investigated the neural mechanisms underlying reactions to protected values in narratives. During fMRI scanning, we presented 78 American, Chinese, and Iranian participants with real-life stories distilled from a corpus of over 20 million weblogs. Reading these stories engaged the posterior medial, medial prefrontal, and temporo-parietal cortices. When participants believed that the protagonist was appealing to a protected value, signal in these regions was increased compared with when no protected value was perceived, possibly reflecting the intensive and iterative search required to process this material. The effect strength also varied across groups, potentially reflecting cultural differences in the degree of concern for protected values.

Laterality of Poststroke Cortical Motor Activity during Action Observation Is Related to Hemispheric Dominance.

Background: Increased activity in the lesioned hemisphere has been related to improved poststroke motor recovery. However, the role of the dominant hemisphere-and its relationship to activity in the lesioned hemisphere-has not been widely explored. Objective: Here, we examined whether the dominant hemisphere drives the lateralization of brain activity after stroke and whether this changes based on if the lesioned hemisphere is the dominant hemisphere or not. Methods: We used fMRI to compare cortical motor activity in the action observation network (AON), motor-related regions that are active both during the observation and execution of an action, in 36 left hemisphere dominant individuals. Twelve individuals had nondominant, right hemisphere stroke, twelve had dominant, left-hemisphere stroke, and twelve were healthy age-matched controls. We previously found that individuals with left dominant stroke show greater ipsilesional activity during action observation. Here, we examined if individuals with nondominant, right hemisphere stroke also showed greater lateralized activity in the ipsilesional, right hemisphere or in the dominant, left hemisphere and compared these results with those of individuals with dominant, left hemisphere stroke. Results: We found that individuals with right hemisphere stroke showed greater activity in the dominant, left hemisphere, rather than the ipsilesional, right hemisphere. This left-lateralized pattern matched that of individuals with left, dominant hemisphere stroke, and both stroke groups differed from the age-matched control group. Conclusions: These findings suggest that action observation is lateralized to the dominant, rather than ipsilesional, hemisphere, which may reflect an interaction between the lesioned hemisphere and the dominant hemisphere in driving lateralization of brain activity after stroke. Hemispheric dominance and laterality should be carefully considered when characterizing poststroke neural activity.

Lesion mapping of cognitive control and value-based decision making in the prefrontal cortex.

A considerable body of previous research on the prefrontal cortex (PFC) has helped characterize the regional specificity of various cognitive functions, such as cognitive control and decision making. Here we provide definitive findings on this topic, using a neuropsychological approach that takes advantage of a unique dataset accrued over several decades. We applied voxel-based lesion-symptom mapping in 344 individuals with focal lesions (165 involving the PFC) who had been tested on a comprehensive battery of neuropsychological tasks. Two distinct functional-anatomical networks were revealed within the PFC: one associated with cognitive control (response inhibition, conflict monitoring, and switching), which included the dorsolateral prefrontal cortex and anterior cingulate cortex and a second associated with value-based decision-making, which included the orbitofrontal, ventromedial, and frontopolar cortex. Furthermore, cognitive control tasks shared a common performance factor related to set shifting that was linked to the rostral anterior cingulate cortex. By contrast, regions in the ventral PFC were required for decision-making. These findings provide detailed causal evidence for a remarkable functional-anatomical specificity in the human PFC.

Persistence of feelings and sentience after bilateral damage of the insula.

It has been convincingly established, over the past decade, that the human insular cortices are involved in processing both body feelings (such as pain) and feelings of emotion. Recently, however, an interpretation of this finding has emerged suggesting that the insular cortices are the necessary and sufficient platform for human feelings, in effect, the sole neural source of feeling experiences. In this study, we investigate this proposal in a patient whose insular cortices were destroyed bilaterally as a result of Herpes simplex encephalitis. The fact that all aspects of feeling were intact indicates that the proposal is problematic. The signals used to assemble the neural substrates of feelings hail from different sectors of the body and are conveyed by neural and humoral pathways to complex and topographically organized nuclei of the brain stem, prior to being conveyed again to cerebral cortices in the somatosensory, insular, and cingulate regions. We suggest that the neural substrate of feeling states is to be found first subcortically and then secondarily repeated at cortical level. The subcortical level would ensure basic feeling states while the cortical level would largely relate feeling states to cognitive processes such as decision-making and imagination.

Sensing, feeling and consciousness.

Living organisms achieve homeostasis by using distinct mechanisms tailored to their physiological complexity. Unicellular organisms as well as plants, which are devoid of nervous systems, rely on covert sensing/detecting and equally covert responding mechanisms. Organisms with nervous systems rely on overt consciousness which is based on homeostatic feelings and the experiences and consequent subjectivity they generate. This article is part of the theme issue 'Sensing and feeling: an integrative approach to sensory processing and emotional experience'.

Correction of eddy current distortions in high angular resolution diffusion imaging.

PURPOSE: To correct distortions caused by eddy currents induced by large diffusion gradients during high angular resolution diffusion imaging without any auxiliary reference scans. MATERIALS AND METHODS: Image distortion parameters were obtained by image coregistration, performed only between diffusion-weighted images with close diffusion gradient orientations. A linear model that describes distortion parameters (translation, scale, and shear) as a function of diffusion gradient directions was numerically computed to allow individualized distortion correction for every diffusion-weighted image. RESULTS: The assumptions of the algorithm were successfully verified in a series of experiments on phantom and human scans. Application of the proposed algorithm in high angular resolution diffusion images markedly reduced eddy current distortions when compared to results obtained with previously published methods. CONCLUSION: The method can correct eddy current artifacts in the high angular resolution diffusion images, and it avoids the problematic procedure of cross-correlating images with significantly different contrasts resulting from very different gradient orientations or strengths.

Understanding otherness: the neural bases of action comprehension and pain empathy in a congenital amputee.

How do we understand and empathize with individuals whose bodies are drastically different from our own? We investigated the neural processes by which an individual with a radically different body, a congenital amputee who is born without limbs, engages her own sensory-motor representations as a means to understand other people's body actions or emotional states. Our results support the prediction that when the goal of the action is possible for the observer, one's own motor regions are involved in processing action observation, just as when individuals viewed those similar to themselves. However, when the observed actions are not possible, mentalizing mechanisms, relying on a different set of neural structures, are additionally recruited to process the actions. Furthermore, our results indicate that when individuals view others experiencing pain in body parts that they have, the insula and somatosensory cortices are activated, consistent with previous reports. However, when an individual views others experiencing pain in body parts that she does not have, the insula and secondary somatosensory cortices are still active, but the primary somatosensory cortices are not. These results provide a novel understanding for how we understand and empathize with individuals who drastically differ from the self.

Hanna and Antonio Damasio.

Hanna and Antonio Damasio work at the intersection between neuroscience, neurology, philosophy, and psychology. They discuss the value of single case studies for neuroscience, consciousness research and the limits of AI, and the fascinating relationship between creativity and the brain.

Seeing touch is correlated with content-specific activity in primary somatosensory cortex.

There is increasing evidence to suggest that primary sensory cortices can become active in the absence of external stimulation in their respective modalities. This occurs, for example, when stimuli processed via one sensory modality imply features characteristic of a different modality; for instance, visual stimuli that imply touch have been observed to activate the primary somatosensory cortex (SI). In the present study, we addressed the question of whether such cross-modal activations are content specific. To this end, we investigated neural activity in the primary somatosensory cortex of subjects who observed human hands engaged in the haptic exploration of different everyday objects. Using multivariate pattern analysis of functional magnetic resonance imaging data, we were able to predict, based exclusively on the activity pattern in SI, which of several objects a subject saw being explored. Along with previous studies that found similar evidence for other modalities, our results suggest that primary sensory cortices represent information relevant for their modality even when this information enters the brain via a different sensory system.

Neural correlates of admiration and compassion.

In an fMRI experiment, participants were exposed to narratives based on true stories designed to evoke admiration and compassion in 4 distinct categories: admiration for virtue (AV), admiration for skill (AS), compassion for social/psychological pain (CSP), and compassion for physical pain (CPP). The goal was to test hypotheses about recruitment of homeostatic, somatosensory, and consciousness-related neural systems during the processing of pain-related (compassion) and non-pain-related (admiration) social emotions along 2 dimensions: emotions about other peoples' social/psychological conditions (AV, CSP) and emotions about others' physical conditions (AS, CPP). Consistent with theoretical accounts, the experience of all 4 emotions engaged brain regions involved in interoceptive representation and homeostatic regulation, including anterior insula, anterior cingulate, hypothalamus, and mesencephalon. However, the study also revealed a previously undescribed pattern within the posteromedial cortices (the ensemble of precuneus, posterior cingulate cortex, and retrosplenial region), an intriguing territory currently known for its involvement in the default mode of brain operation and in self-related/consciousness processes: emotions pertaining to social/psychological and physical situations engaged different networks aligned, respectively, with interoceptive and exteroceptive neural systems. Finally, within the anterior insula, activity correlated with AV and CSP peaked later and was more sustained than that associated with CPP. Our findings contribute insights on the functions of the posteromedial cortices and on the recruitment of the anterior insula in social emotions concerned with physical versus psychological pain.

A hybrid high-resolution anatomical MRI atlas with sub-parcellation of cortical gyri using resting fMRI.

We present a new high-quality, single-subject atlas with sub-millimeter voxel resolution, high SNR, and excellent gray-white tissue contrast to resolve fine anatomical details. The atlas is labeled into two parcellation schemes: 1) the anatomical BCI-DNI atlas, which is manually labeled based on known morphological and anatomical features, and 2) the hybrid USCBrain atlas, which incorporates functional information to guide the sub-parcellation of cerebral cortex. In both cases, we provide consistent volumetric and cortical surface-based parcellation and labeling. The intended use of the atlas is as a reference template for structural coregistration and labeling of individual brains. A single-subject T1-weighted image was acquired five times at a resolution of 0.547 mm × 0.547 mm × 0.800 mm and averaged. Images were processed by an expert neuroanatomist using semi-automated methods in BrainSuite to extract the brain, classify tissue-types, and render anatomical surfaces. Sixty-six cortical and 29 noncortical regions were manually labeled to generate the BCI-DNI atlas. The cortical regions were further sub-parcellated into 130 cortical regions based on multi-subject connectivity analysis using resting fMRI (rfMRI) data from the Human Connectome Project (HCP) database to produce the USCBrain atlas. In addition, we provide a delineation between sulcal valleys and gyral crowns, which offer an additional set of 26 sulcal subregions per hemisphere. Lastly, a probabilistic map is provided to give users a quantitative measure of reliability for each gyral subdivision. Utility of the atlas was assessed by computing Adjusted Rand Indices (ARIs) between individual sub-parcellations obtained through structural-only coregistration to the USCBrain atlas and sub-parcellations obtained directly from each subject's resting fMRI data. Both atlas parcellations can be used with the BrainSuite, FreeSurfer, and FSL software packages.

Comparison of landmark-based and automatic methods for cortical surface registration.

Group analysis of structure or function in cerebral cortex typically involves, as a first step, the alignment of cortices. A surface-based approach to this problem treats the cortex as a convoluted surface and coregisters across subjects so that cortical landmarks or features are aligned. This registration can be performed using curves representing sulcal fundi and gyral crowns to constrain the mapping. Alternatively, registration can be based on the alignment of curvature metrics computed over the entire cortical surface. The former approach typically involves some degree of user interaction in defining the sulcal and gyral landmarks while the latter methods can be completely automated. Here we introduce a cortical delineation protocol consisting of 26 consistent landmarks spanning the entire cortical surface. We then compare the performance of a landmark-based registration method that uses this protocol with that of two automatic methods implemented in the software packages FreeSurfer and BrainVoyager. We compare performance in terms of discrepancy maps between the different methods, the accuracy with which regions of interest are aligned, and the ability of the automated methods to correctly align standard cortical landmarks. Our results show similar performance for ROIs in the perisylvian region for the landmark-based method and FreeSurfer. However, the discrepancy maps showed larger variability between methods in occipital and frontal cortex and automated methods often produce misalignment of standard cortical landmarks. Consequently, selection of the registration approach should consider the importance of accurate sulcal alignment for the specific task for which coregistration is being performed. When automatic methods are used, the users should ensure that sulci in regions of interest in their studies are adequately aligned before proceeding with subsequent analysis.

Sulcal set optimization for cortical surface registration.

Flat mapping based cortical surface registration constrained by manually traced sulcal curves has been widely used for inter subject comparisons of neuroanatomical data. Even for an experienced neuroanatomist, manual sulcal tracing can be quite time consuming, with the cost increasing with the number of sulcal curves used for registration. We present a method for estimation of an optimal subset of size N(C) from N possible candidate sulcal curves that minimizes a mean squared error metric over all combinations of N(C) curves. The resulting procedure allows us to estimate a subset with a reduced number of curves to be traced as part of the registration procedure leading to optimal use of manual labeling effort for registration. To minimize the error metric we analyze the correlation structure of the errors in the sulcal curves by modeling them as a multivariate Gaussian distribution. For a given subset of sulci used as constraints in surface registration, the proposed model estimates registration error based on the correlation structure of the sulcal errors. The optimal subset of constraint curves consists of the N(C) sulci that jointly minimize the estimated error variance for the subset of unconstrained curves conditioned on the N(C) constraint curves. The optimal subsets of sulci are presented and the estimated and actual registration errors for these subsets are computed.

Changes in auditory cortical thickness following music training in children: converging longitudinal and cross-sectional results.

Evidence is accumulating to suggest that music training is associated with structural brain differences in children and in adults. We used magnetic resonance imagining in two studies to investigate neuroanatomical correlates of music training in children. In study 1, we cross-sectionally compared a group of child musician (ages 9-11) matched to non-musicians and found that cortical thickness was greater in child musician in the posterior segment of the right-superior temporal gyrus (STG), an auditory association area that is involved in processing complex auditory stimuli, including pitch. We also found that thickness in the right posterior STG is related to music proficiency, however this relationship did not reach significance. In study 2, a longitudinal study, we investigated change in cortical thickness over a four-year period comparing a group of children involved in a systematic music training program with another group of children who did not have any music training. In this 2nd study we assessed both groups at the beginning of the study, prior to music training for the music group, and four years later. We found that children in the music group showed a strong trend of lower rate of cortical thinning in the right posterior superior temporal gyrus. Together, our results provide evidence that music training induces structural brain changes in school-age children and that these changes are predominantly pronounced in the right auditory association areas.

Morphology of the insula in relation to hearing status and sign language experience.

We investigated whether auditory deprivation and/or sign language exposure during development alters the macroscopic neuroanatomy of the human insula. Volumetric analyses were based on MRI data from 25 congenitally deaf subjects who were native users of American Sign Language (ASL), 25 hearing subjects with no knowledge of ASL, and 16 hearing subjects who grew up in deaf families and were native ASL signers. Significant variation in insula volume was associated with both hearing status and sign language experience. Compared with both hearing groups, deaf subjects exhibited a significant increase in the amount of gray matter in the left posterior insular lobule, which we hypothesize may be related to the dependence on lip-reading and articulatory-based (rather than auditory-based) representations of speech for deaf individuals. Both deaf and hearing signers exhibited an increased volume of white matter in the right insula compared with hearing nonsigners. We hypothesize that the distinct morphology of the right insula for ASL signers may arise from enhanced connectivity resulting from an increased reliance on cross-modal sensory integration in sign language compared with spoken language.