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Is cortex necessary?
... However, this type of evidence faces problems that parallel those discussed above. First, while Långsjö and colleagues' results highlight the importance of the midbrain in restoring consciousness, they also highlight the importance of the anterior cingulate cortex (ACC) and connections between the ACC and parietal and frontal cortical regions (a criticism raised by Allen-Hermanson, 2016). They describe this as "only minimal cortical activity", and Klein and Barron (2016b, pp. ...
There is no agreement on whether any invertebrates are conscious and no agreement on a methodology that could settle the issue. How can the debate move forward? I distinguish three broad types of approach: theory-heavy, theory-neutral and theory-light. Theory-heavy and theory-neutral approaches face serious problems, motivating a middle path: the theory-light approach. At the core of the theory-light approach is a minimal commitment about the relation between phenomenal consciousness and cognition that is compatible with many specific theories of consciousness: the hypothesis that phenomenally conscious perception of a stimulus facilitates, relative to unconscious perception, a cluster of cognitive abilities in relation to that stimulus. This “facilitation hypothesis” can productively guide inquiry into invertebrate consciousness. What is needed? At this stage, not more theory, and not more undirected data gathering. What is needed is a systematic search for consciousness-linked cognitive abilities, their relationships to each other, and their sensitivity to masking.
The entomology literature has historically suggested insects cannot feel pain, leading to their exclusion from ethical debates and animal welfare legislation. However, there may be more neural and cognitive/behavioural evidence for pain in insects than previously considered. We use Birch et al. 's (2021) eight criteria for sentience to critically evaluate the evidence for pain in insects. We assess six orders (Blattodea, Coleoptera, Diptera, Hymenoptera, Lepidoptera, and Orthoptera) in at least two life stages (adult and first instar juveniles, as well as other instars where relevant data are found). Other insect orders have not received enough research effort to be evaluated. According to the Birch et al. framework, adult Diptera (flies and mosquitoes) and Blattodea (cockroaches and termites) satisfy six criteria, constituting strong evidence for pain. Adults of the remaining orders (except Coleoptera, beetles) and some juveniles (Blattodea and Diptera, as well as last instar Lepidoptera [butterflies and moths]) satisfy 3–4 criteria, or “substantial evidence for pain”. We found no good evidence that any insects failed a criterion. However, there were significant evidence gaps, particularly for juveniles, highlighting the importance of more research on insect pain. We conclude by considering the ethical implications of our findings where insects are managed in wild, farmed, and research contexts.
How subjective experience is realized in nervous systems remains one of the great challenges in the natural sciences. An answer to this question should resolve debate about which animals are capable of subjective experience. We contend that subjective experience of sensory stimuli is dependent on the brain’s awareness of its internal neural processing of these stimuli. This premise is supported by empirical evidence demonstrating that disruption to either processing streams or awareness states perturb subjective experience. Given that the brain must predict the nature of sensory stimuli, we reason that conscious awareness is itself dependent on predictions generated by hierarchically organized forward models of the organism’s internal sensory processing. The operation of these forward models requires a specialized neural architecture and hence any nervous system lacking this architecture is unable to subjectively experience sensory stimuli. This approach removes difficulties associated with extrapolations from behavioral and brain homologies typically employed in addressing whether an animal can feel. Using nociception as a model sensation, we show here that the Drosophila brain lacks the required internal neural connectivity to implement the computations required of hierarchical forward models. Consequently, we conclude that Drosophila, and those insects with similar neuroanatomy, do not subjectively experience noxious stimuli and therefore cannot feel pain.
To what degree are non-human animals conscious? We propose that the most meaningful way to approach this question is from the perspective of functional neurobiology. Here we focus on subjective experience, which is a basic awareness of the world without further reflection on that awareness. This is considered the most basic form of consciousness. Tellingly, this capacity is supported by the integrated midbrain and basal ganglia structures, which are among the oldest and most highly conserved brain systems in vertebrates. A reasonable inference is that the capacity for subjective experience is both widespread and evolutionarily old within the vertebrate lineage. We argue that the insect brain supports functions analogous to those of the vertebrate midbrain and hence that insects may also have a capacity for subjective experience. We discuss the features of neural systems which can and cannot be expected to support this capacity as well as the relationship between our arguments based on neurobiological mechanism and our approach to the " hard problem " of conscious experience.
How, why, and when consciousness evolved remain hotly debated topics. Addressing these issues requires considering the distribution of consciousness across the animal phylogenetic tree. Here we propose that at least one invertebrate clade, the insects, has a capacity for the most basic aspect of consciousness: subjective experience. In vertebrates the capacity for subjective experience is supported by integrated structures in the midbrain that create a neural simulation of the state of the mobile animal in space. This integrated and egocentric representation of the world from the animal's perspective is sufficient for subjective experience. Structures in the insect brain perform analogous functions. Therefore, we argue the insect brain also supports a capacity for subjective experience. In both vertebrates and insects this form of behavioral control system evolved as an efficient solution to basic problems of sensory reafference and true navigation. The brain structures that support subjective experience in vertebrates and insects are very different from each other, but in both cases they are basal to each clade. Hence we propose the origins of subjective experience can be traced to the Cambrian.
Are animals conscious? If so, when did consciousness evolve? We address these long-standing and essential questions using a modern neuroscientific approach that draws on diverse fields such as consciousness studies, evolutionary neurobiology, animal psychology, and anesthesiology. We propose that the stepwise emergence from general anesthesia can serve as a reproducible model to study the evolution of consciousness across various species and use current data from anesthesiology to shed light on the phylogeny of consciousness. Ultimately, we conclude that the neurobiological structure of the vertebrate central nervous system is evolutionarily ancient and highly conserved across species and that the basic neurophysiologic mechanisms supporting consciousness in humans are found at the earliest points of vertebrate brain evolution. Thus, in agreement with Darwin's insight and the recent "Cambridge Declaration on Consciousness in Non-Human Animals," a review of modern scientific data suggests that the differences between species in terms of the ability to experience the world is one of degree and not kind.
Feelings are mental experiences of body states. They signify physiological need (for example, hunger), tissue injury (for example, pain), optimal function (for example, well-being), threats to the organism (for example, fear or anger) or specific social interactions (for example, compassion, gratitude or love). Feelings constitute a crucial component of the mechanisms of life regulation, from simple to complex. Their neural substrates can be found at all levels of the nervous system, from individual neurons to subcortical nuclei and cortical regions.
It has been proposed that self-awareness (SA), a multifaceted phenomenon central to human consciousness, depends critically on specific brain regions, namely the insular cortex, the anterior cingulate cortex (ACC), and the medial prefrontal cortex (mPFC). Such a proposal predicts that damage to these regions should disrupt or even abolish SA. We tested this prediction in a rare neurological patient with extensive bilateral brain damage encompassing the insula, ACC, mPFC, and the medial temporal lobes. In spite of severe amnesia, which partially affected his "autobiographical self", the patient's SA remained fundamentally intact. His Core SA, including basic self-recognition and sense of self-agency, was preserved. His Extended SA and Introspective SA were also largely intact, as he has a stable self-concept and intact higher-order metacognitive abilities. The results suggest that the insular cortex, ACC and mPFC are not required for most aspects of SA. Our findings are compatible with the hypothesis that SA is likely to emerge from more distributed interactions among brain networks including those in the brainstem, thalamus, and posteromedial cortices.
One of the greatest challenges of modern neuroscience is to discover the neural mechanisms of consciousness and to explain how they produce the conscious state. We sought the underlying neural substrate of human consciousness by manipulating the level of consciousness in volunteers with anesthetic agents and visualizing the resultant changes in brain activity using regional cerebral blood flow imaging with positron emission tomography. Study design and methodology were chosen to dissociate the state-related changes in consciousness from the effects of the anesthetic drugs. We found the emergence of consciousness, as assessed with a motor response to a spoken command, to be associated with the activation of a core network involving subcortical and limbic regions that become functionally coupled with parts of frontal and inferior parietal cortices upon awakening from unconsciousness. The neural core of consciousness thus involves forebrain arousal acting to link motor intentions originating in posterior sensory integration regions with motor action control arising in more anterior brain regions. These findings reveal the clearest picture yet of the minimal neural correlates required for a conscious state to emerge.
A compelling single case report of visual awareness (visual qualia) without primary visual cortex would be sufficient to refute the hypothesis that the primary visual cortex and the back-projections to it are necessary for conscious visual experience. In a previous study, we emphasized the presence of crude visual awareness in Patient G.Y., with a lesion of the primary visual cortex, who is aware of, and able to discriminate, fast-moving visual stimuli presented to his blind field. The visual nature of Patient G.Y.'s blind field experience has since been questioned and it has been suggested that the special circumstances of repeated testing over decades may have altered Patient G.Y.'s visual pathways. We therefore sought new evidence of visual awareness without primary visual cortex in patients for whom such considerations do not apply. Three patients with hemianopic field defects (Patient G.N. and Patient F.B. with MRI confirmed primary visual cortex lesions, Patient C.G. with an inferred lesion) underwent detailed psychophysical testing in their blind fields. Visual stimuli were presented at different velocities and contrasts in two- and four-direction discrimination experiments and the direction of motion and awareness reported using a forced-choice paradigm. Detailed verbal reports were also obtained of the nature of the blind field experience with comparison of the drawings of the stimulus presented in the blind and intact fields, where possible. All three patients reported visual awareness in their blind fields. Visual awareness was significantly more likely when a moving stimulus was present compared to no stimulus catch trials (P < 0.01 for each subject). Psychophysical performance in Patient F.B. and Patient G.N. was consistent with the Riddoch syndrome, with higher levels of visual awareness for moving compared to static stimuli (P < 0.001) and intact direction discrimination (P < 0.0001 for two- and four-direction experiments). Although the blind field experience of all three subjects was degraded, it was clearly visual in nature. We conclude that the primary visual cortex or back-projections to it are not necessary for visual awareness.
Despite converging agreement about the definition of persistent vegetative state, recent reports have raised concerns about the accuracy of diagnosis in some patients, and the extent to which, in a selection of cases, residual cognitive functions may remain undetected. Objective assessment of residual cognitive function can be extremely difficult as motor responses may be minimal, inconsistent, and difficult to document in many patients, or may be undetectable in others because no cognitive output is possible. Here we describe strategies for using H(2)(15)O positron emission tomography activation studies to study covert cognitive processing in patients with a clinical diagnosis of persistent vegetative state. Three cases are described in detail. Of these, two exhibited clear and predicted regional cerebral blood flow responses during well-documented activation paradigms (face recognition and speech perception) which have been shown to produce specific, robust and reproducible activation patterns in normal volunteers. Some months after scanning, both patients made a significant recovery. In a third case, blood flow data were acquired during a speech perception task, although methodological difficulties precluded any systematic interpretation of the results. In spite of the multiple logistic and procedural problems involved, these results have major clinical and scientific implications and provide a strong basis for the systematic study of possible residual cognitive function in patients diagnosed as being in a persistent vegetative state.
Although lesions of the striate (V1) cortex disrupt conscious vision, patients can demonstrate surprising residual abilities within their affected visual field, a phenomenon termed blindsight. The relative contribution of spared "islands" of functioning striate cortex to residual vision, versus subcortical pathways to extrastriate areas, has implications for the role of early visual areas in visual awareness and performance. Here we describe the behavioral and neural features of residual cortical function in Patient M.C., who sustained a posterior cerebral artery stroke at the age of 15 years. Within her impaired visual field, we found preserved visual abilities characteristic of blindsight, including superior detection of motion, and above-chance discrimination of shape, color, and motion direction. Functional magnetic resonance imaging demonstrated a retinotopically organized representation of M.C.'s blind visual field within the lesioned occipital lobe, specifically within area V1. The incongruity of a well-organized cortex and M.C.'s markedly impaired vision was resolved by measurement of functional responses within her damaged occipital lobe. Attenuated neural contrast-response functions were found to correlate with M.C.'s impaired psychophysical performance. These results demonstrate that the behavioral features of blindsight may arise in the presence of residual striate responses that are spatially organized and sensitive to contrast variation.
AimHydranencephaly is commonly taken to exemplify the developmental vegetative state, based largely on the assumption that radical loss of cortical tissue is incompatible with consciousness. The aim of the study reported here was to survey primary caregivers of children born with hydranencephaly for behavioural evidence indirectly informative about the conscious status of these children. Methods
The survey recruited 108 primary caregivers through a parent support group and was conducted online via a commercial survey hosting facility. As part of a more extensive questionnaire, participants answered 106 questions bearing on the environmental responsiveness, emotional reactivity, mood and agency of the child in their care. ResultsThe survey elicited a many-facetted and detailed set of caregiver answers and written observations regarding the child's behaviour. A conservative measure of agreement among respondents' answers yielded a generic portrait of the responsiveness and expressive behaviour of a hydranencephaly child. Conclusion
The generic behavioural characteristics of hydranencephaly thus assessed are incompatible on multiple counts with the unconsciousness characteristic of the vegetative state. This bears on what is included under the concept of quality of life for children with hydranencephaly, and hence on appropriate forms of treatment and care in their case.
Neurophysiological and neuroimaging studies including both patients with disorders of consciousness and healthy subjects with modified states of consciousness suggest a crucial role of the medial posteroparietal cortex in conscious information processing. However no direct neuropsychological evidence supports this hypothesis and studies including patients with restricted lesions of this brain region are almost non-existent. Using direct intraoperative electrostimulations, we showed in a rare patient that disrupting the subcortical connectivity of the left posterior cingulate cortex (PCC) reliably induced a breakdown in conscious experience. This acute phenomenon was mainly characterized by a transient behavioral unresponsiveness with loss of external connectedness. In all cases, when he regained consciousness, the patient described himself as in dream, outside the operating room. This finding suggests that functional integrity of the PPC connectivity is necessary for maintaining consciousness of external environment.
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.
Recovery of consciousness following severe brain injuries can occur over long time intervals. Importantly, evolving cognitive recovery can be strongly dissociated from motor recovery in some individuals, resulting in underestimation of cognitive capacities. Common mechanisms of cerebral dysfunction that arise at the neuronal population level may explain slow functional recoveries from severe brain injuries. This review proposes a "mesocircuit" model that predicts specific roles for different structural and dynamic changes that may occur gradually during recovery. Recent functional neuroimaging studies that operationally identify varying levels of awareness, memory and other higher brain functions in patients with no behavioral evidence of these cognitive capacities are discussed. Measuring evolving changes in underlying brain function and dynamics post-injury and post-treatment frames future investigative work.
The minimally conscious state (MCS) resulting from severe brain damage refers to a subset of patients who demonstrate unequivocal, but intermittent, behavioral evidence of awareness of self or their environment. Although clinical examination may suggest residual cognitive function, neurobiological correlates of putative cognition in MCS have not been demonstrated.
To test the hypothesis that MCS patients retain active cerebral networks that underlie cognitive function even though command following and communication abilities are inconsistent.
fMRI was employed to investigate cortical responses to passive language and tactile stimulation in two male adults with severe brain injuries leading to MCS and in seven healthy volunteers.
In the case of the patient language-related tasks, auditory stimulation with personalized narratives elicited cortical activity in the superior and middle temporal gyrus. The healthy volunteers imaged during comparable passive language stimulation demonstrated responses similar to the patients' responses. However, when the narratives were presented as a time-reversed signal, and therefore without linguistic content, the MCS patients demonstrated markedly reduced responses as compared with volunteer subjects, suggesting reduced engagement for "linguistically" meaningless stimuli.
The first fMRI maps of cortical activity associated with language processing and tactile stimulation of patients in the minimally conscious state (MCS) are presented. These findings of active cortical networks that serve language functions suggest that some MCS patients may retain widely distributed cortical systems with potential for cognitive and sensory function despite their inability to follow simple instructions or communicate reliably.
The issue of the biological origin of consciousness is linked to that of its function. One source of evidence in this regard is the contrast between the types of information that are and are not included within its compass. Consciousness presents us with a stable arena for our actions-the world-but excludes awareness of the multiple sensory and sensorimotor transformations through which the image of that world is extracted from the confounding influence of self-produced motion of multiple receptor arrays mounted on multijointed and swivelling body parts. Likewise excluded are the complex orchestrations of thousands of muscle movements routinely involved in the pursuit of our goals. This suggests that consciousness arose as a solution to problems in the logistics of decision making in mobile animals with centralized brains, and has correspondingly ancient roots.
A broad range of evidence regarding the functional organization of the vertebrate brain - spanning from comparative neurology to experimental psychology and neurophysiology to clinical data - is reviewed for its bearing on conceptions of the neural organization of consciousness. A novel principle relating target selection, action selection, and motivation to one another, as a means to optimize integration for action in real time, is introduced. With its help, the principal macrosystems of the vertebrate brain can be seen to form a centralized functional design in which an upper brain stem system organized for conscious function performs a penultimate step in action control. This upper brain stem system retained a key role throughout the evolutionary process by which an expanding forebrain - culminating in the cerebral cortex of mammals - came to serve as a medium for the elaboration of conscious contents. This highly conserved upper brainstem system, which extends from the roof of the midbrain to the basal diencephalon, integrates the massively parallel and distributed information capacity of the cerebral hemispheres into the limited-capacity, sequential mode of operation required for coherent behavior. It maintains special connective relations with cortical territories implicated in attentional and conscious functions, but is not rendered nonfunctional in the absence of cortical input. This helps explain the purposive, goal-directed behavior exhibited by mammals after experimental decortication, as well as the evidence that children born without a cortex are conscious. Taken together these circumstances suggest that brainstem mechanisms are integral to the constitution of the conscious state, and that an adequate account of neural mechanisms of conscious function cannot be confined to the thalamocortical complex alone.
Hydranencephaly is a rare neurological condition in which the cerebral hemispheres are either absent or severely compromised. It is widely believed that children with hydranencephaly are not conscious; and therefore, are routinely classified into the diagnostic criteria of vegetative state. However, there are several pieces of behavioral evidence clearly indicating the presence of consciousness in such patients. Here, I review these behavioral evidence and argue how misclassification of these patients and assigning them a lack of consciousness have far-reaching implications in terms of both clinical and theoretical neuroscience.