Mental Radio (Studies in Consciousness)

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Given the widely held causal relation between brain and behavior, an important question for psychology, neuroscience and even philosophy is whether and to what extent all the different vertebrate species share the mechanisms involved in the evolutionary emergence of phenomenal experiences including affective, interoceptive, and exteroceptive varieties to the much higher ability to mentally represent the self as an actor in the world.

In the following we examine this extant issue by starting from humans in whom a variety of simple and complex forms of consciousness exist. Knowledge about the meaning of world and self-representation deriving from phenomenological, neuropsychological and neurophysiological approaches will be reviewed. Since phenomenological subjective evidence can be obtained only in humans but only semi-directly through language , we must tackle evolutionarily related issues in the other classes of vertebrates fishes, amphibians, reptiles, birds, and mammals by reviewing not only neuroanatomical, neuropsychological and neurophysiological data.

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Moreover behavioral data hinting at the presence of affective experiences will be discussed by evaluating the rewarding and punishing properties, as inferred from deep brain stimulation DBS of brain emotional circuits. It is important to note that we do not intend to use the phylogenetic taxonomy for establishing a sort of hierarchy among vertebrates and we do not intend to attribute to, say, living reptiles a higher or lower place with respect to mammals. We simply mean that the different classes of vertebrates represent different genealogical lines originating from a common origin and that each class can be described by referring to the complexity of their nervous system Butler and Hodos, ; Denton, , and to functional homologies of the brain structures involved in primary-process emotions Panksepp, a , In the following, we discuss homologies, analogies and differences in the way self and world representation manifest in the different vertebrate species.

Neuroscience studies indicate that the human brain continuously creates the conscious experience of being in the world i. Consciousness emerges from the brain's ability to construct a complex representation of the world and the self Edelman, That both the self and the world are constructed by the brain is strongly suggested by the profound changes of self and world consciousness following cerebral lesions and during dreaming, a condition where external inputs are blocked Frith, It is worth noting that lesions of specific brain regions may bring about domain-specific deficits of awareness.

Lesions to different visual areas, for example, may induce defects in the recognition the form apperceptive agnosia or the movement of an object akinetopsia, Zeki, and lesions to higher-order cortical regions may induce defective awareness of space e. Complex domain-specific deficits following frontal or occipito-parietal regions such as anosoagnosia for hemiplegia Moro et al. However, we are not aware that our experience of the world and of the self are brain constructions and that self and world represent models through which information is processed.

In philosophical parlance, the illusion that such models are the reality is called transparency Revonsuo, ; Metzinger, It is worth noting that the brain's construction of the world and self is not arbitrary but it takes into account the operational interactions learned at both phylogenetic and ontogenetic levels Jerison, ; Striedter, More specifically, the world representations created by the brain are highly adaptive and allow individuals to implement a number of effective operations like moving in the world, maintaining bodily and affective homeostasis, developing plans for self- fulfillment Geary, Conscious awareness is a simulation per se and does not put the subject in direct contact with reality.

It is thus possible to think of mental experiments in which even an expert pilot unaware to be in a very realistic flight simulator believes he is operating on a real airplane Metzinger, Moreover, representations concerning the world and the self may often be contradictory, as indicated by the fact that illusions may fool vision but not action suggesting errors may occur in allocentric i. Crucially, the same participants who are fooled by the illusion when making perceptual judgments, when asked to pick up the central circle scale their grip aperture on the basis of the true size of the target disc and not of its illusory size Aglioti et al.

In a complementary vein, illusory doubling of tactile stimuli may occur only if a conflict occurs between egocentric body-based and allocentric world centered coordinate frames like for example in the Aristotle's diploaesthesia. Indeed, one object touching the contact area between the crossed index and middle fingers is perceived as two objects localized on the lateral surface of each finger Bufalari et al. Striking contradictions occur also during dreaming Hobson, , , suggesting an analogy between this state of consciousness and the above illusions. All these states may be of fundamental importance for understanding the problem of consciousness Metzinger, ; Revonsuo, Human consciousness can be analyzed according to different levels, the most significant of which are the phenomenological, the neurological and the neurophysiological ones Revonsuo, Each level possesses specific properties and rules.

We acknowledge, however, that different epistemological perspectives suggest different models ranging from hierarchical Craver and Darden, ; Neisser, , to relatively independent ones Ayala and Arp, In addition to the classical third-person-perspective analysis 3PP: a perspective in which the whole universe is given as existent independently from the observer it is necessary to envisage a first person perspective 1PP: in this perspective the objects in space are necessarily related to an agent Revonsuo, , At this level, consciousness is an immediate, undeniable fact of experience, i.

The ego-center is single and metaphorically located behind the bridge of the nose, inside our head Merker, , but more realistically arising from certain within-brain experiences. The objects belonging to the world are located in the phenomenal dimension characterized by spatial and temporal attributes Revonsuo, Already at the phenomenological level, the self does not seem to be an object, rather a within-brain process, a continuous stream of experiences and thoughts.

The self seems to disappear during deep slow-wave sleep, and sometimes to dissolve or have less defined boundaries, also during dreaming Revonsuo, Tellingly, the self reappears with greater perceptual depth and clarity when we wake up, as already noticed by the Heraclitus more than 25 centuries ago Haxton and Hillman, The self seems to consist of at least two components, probably arranged in layers.

An exception to this neglect may come from psychoanalysis that made serious attempts to distinguish between the Ego and the Self Treurniet, Human consciousness has been operationalized as mainly formed by two components, namely arousal wakefulness or vigilance, with phenomenal contents and awareness which implies higher recognition of those contents.

This latter component is further divided in external awareness concerning the sensorial analysis of the environment and the internal awareness or self-consciousness concerning the inner mental representation of the self Demertzi et al. While arousal is linked to activity of brain stem neural populations connected both directly or indirectly with the cerebral cortex, awareness depends on the functional integrity of cerebral cortex and thalamus.

From an evolutionary perspective, the former is commonly assumed to be foundational for the latter. Neuroimaging studies indicate that awareness depends on the integrity of a large cortical network with medial fronto-parietal structures dealing with processing of internal states mainly related to self-consciousness mind wandering, inner speech, autobiographical memory recall and lateral cortical structures mainly dealing with awareness of the external world Vanhaudenhuyse et al.

Fundamental clues to the understanding of structures and mechanisms necessary and sufficient to the appearance of conscious self and world representation come from the analysis of patients in whom cerebral damage alters different forms of consciousness, like coma, vegetative state VS , and minimally conscious state MCS Laureys et al.

Functional imaging studies indicate that medial and posterior cortical regions like the precuneus and the posterior cingulate cortex play a crucial role in human awareness. These regions, for example, do show high metabolic activity during aware wakefulness. By contrast they are deactivated in profound anesthesia, in VS patients and in severely demented patients.

Relevant to this issue are the recent studies using transcranial magnetic stimulation TMS to perturb the cortex and EEG to record the effects of such perturbation. These studies demonstrate the importance of cortico-cortical and cortico-subcortical functional connectivity in human conscious awareness. More specifically the perturbation induced by TMS on EEG in VS patients indicate a severe defect of connectivity that is reminiscent of deep sleep and general anaesthesia states Casali et al.

Such complex EEG activity, typically found in in healthy controls, is also observed in Locked-in patients who are aware of the self and of the environment in spite of their severe de-efferentation Rosanova et al.

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The notion that consciousness is related to an optimal functional connectivity is in keeping with influential theories of consciousness developed in psychology e. Empirical and philosophical studies suggest that the self, in human beings, is formed by diverse neuropsychological components Stern, ; Gallagher, ; Northoff et al. This distinction reappeared in the contemporary neuroscience debate. Particularly important are the studies of complex human pathologies, e. Overall the following components of human self can be defined: i the proto-self; ii the core self; iii the self-consciousness; iv the narrative self.

It can be considered as an organizational unit providing some kind of foundational coordination for all sensations Humphrey, , and primordial emotions Panksepp, a , b ; Denton, Behavioral studies indicate that human newborns do exhibit a complex affective life as indexed by basic emotions as pain, joy, disgust, anger and by self-relevant acts like hand-mouth coordination movements Stern, ; Rochat, For example, newborn babies may purposely move their hand to counteract external forces applied to their wrist with the clear aim to keep seeing their hand and thus exert a better control on their own action Van der Meer and Lee, It is also relevant that human neonates exhibit a higher number of head turns toward tactile stimuli delivered on their cheek by an examiner external stimulation with respect to tactile stimuli delivered on the cheek by themselves self-stimulation.

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This behavioral pattern speaks in favor of the innate ability to discriminate whether tactile stimuli are delivered by the self vs. The core self involves a number of components, including the sense of ownership i. These dimensions are pre-reflective, implicit, or tacit aspects of our experience Vogeley and Gallagher, , p. The core self is connected to semantic memory systems and also to affective systems Tulving, a ; Northoff and Panksepp, ; Panksepp and Northoff, Such basic forms of self and world representations likely operate only in the present moment Edelman, ; Edelman and Tononi, Individuals who possess this form of consciousness show self-recognition and recognition of objects in the world Tulving, b.

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Many nonhuman animals, especially mammals and birds, for example, seem to be endowed with well-developed knowledge-of-the-world semantic memory and to flexibly utilize this information Tulving, a , p. Conditioning studies indicate that semantic memory systems are present in birds. Moreover, selective lesions of the teleost lateral pallium a structure analogous to the mammalian hippocampus in goldfish trained in a variety of spatial memory tasks induce conspicuous spatial memory deficits Broglio et al.

Two important characteristics of the core self are the sense of ownership i. Only at the age of 18 months children become capable of self-recognizing themselves in the mirror and to understand that pictures represent other people. According to Perner mirror self-recognition implies to own the representation of the model of a real self as well as the model of the reflected-in-a-mirror model.

The same test has been used in several vertebrate species. Although controversies have arisen about the validity of the mark test, studies suggest that chimpanzees after the age of 28 month , orangutans, macaques, dolphins, elephants and corvid birds may show evidence of mirror self-exploration.

It is worth noting that the mirror test may not represent definitive evidence for the presence or absence of self-recognition and spurious variables like scarce motivation to watch reflected images may influence the test Gallup et al. The narrative self Gallagher, ; Boyd, ; Damasio, refers to the capability of handle episodic-type, declarative memories that unify the self into a coherent story.

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Prerequisites for developing a fully blown narrative self may include language Gazzaniga, , ; Baddeley et al. Such a device makes possible not only the aware recall of past events i. The ability to mental traveling in time seems to be underpinned by a variety of neural regions involved in memory e. The capability to mentally travel in time, an ability linked to the task-unrelated thoughts that characterize mind wandering, probably appeared more than one million years ago in the genus Homo habilis and is possibly at the basis of building lithic tools Corballis, ; Fabbro and Crescentini, Importantly, the mind may wander not only in time by also in others' mind.

Related to this is the development of shared intentionality joint intentions and attention and the ability to know or the belief to be able to know what other individuals think or believe, the so-called theory of mind that seems to be mainly a human feature Tomasello et al. It has been suggested that self-referential cognition, inner speech, mind wandering and autobiographical memory are important components of self-awareness a complex function likely underpinned by medial brain areas Vanhaudenhuyse et al.

One of the most significant discoveries in the contemporary neurophysiological research has been the correlation between neural activity of specific neuronal populations and conscious experience.

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It has been demonstrated that the conscious recognition of a visual stimulus correlates with synchronized discharge frequencies of neurons around 40 Hz or in the gamma band: 20—80 Hz. In binocular rivalry experiments Logothetis and Schall, , for example, the gamma band is correlated with the activity of the visual areas that are involved in the processing of the dominant conscious stimulus. The same gamma band activity decreases in neurons that are involved in the processing of suppressed non-conscious stimuli Engel and Singer, ; Fries, Interestingly, such activity has been found also at the subcortical level e.

Neurophysiology may help substantially to understand the link between states of consciousness and brain activity.

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Seminal studies indicate that it takes a long time about — ms to sensory stimuli to reach the threshold of conscious awareness Libet et al. PCI discriminated different levels of consciousness not only in healthy individuals during wakefulness, sleep, and anesthesia, but also in patients who had emerged from coma and recovered a minimal level of consciousness. In particular, inter-cortical connectivity was higher during awake and REM states and lower during slow-waves sleep, anaesthesia and minimal consciousness states Casali et al.

Artistic representation of the basal and forebrain regions in the vertebrates Image credits to Massimo Bergamasco. Lateral view Left panel and mid-sagittal view right panel of the brain of: A rainbow trout, Salmo gairdneri; B green frog, Rana esculenta; C tegu lizard, Tupinambis teguixin; D pigeon, Columba livia; E cat; and F human brain.