Mauro, the question of creativity, like that of consciousness, is a difficult and challenging one. Sure, fMRI can detect electrical changes in parts of the brain consistent with the creative process, but are they causal? or merely correlations and associations? I think that creativity is much more fundamental than just brain electrical activity. I think it started at the beginning of life itself, demarcating the external and internal 'environments'. That process is recapitulated each time the life cycle begins, sperm penetrating egg, causing a 'spark' of electrical activity that persists throughout the life of the organism until it dies. It is the foundation of embryologic development and growth, forming the cellular interactions that generate the organism. The homeostatic and allostatic regulatory mechanisms that determine the electrical activity of all of the organs, including the brain are the result of physiologic development that has evolved from the unicellular state and returns to it iteratively during reproduction. The creative 'spark' recapitulates those processes, and is at its maximum when it resonates throughout both the body and mind, what Mazlow referred to as 'peak moments'. As for the mechanisms of communication between the body and brain, we know much more about the signals from the brain to the body, but there are those that travel from the body to the brain. The big breakthrough in the study of brain evolutoin was the realization (Nick Holland) that the invertebrate central nervous system is in the skin, giving rise to the concept of 'skin-brain'. That concept offers insights into the role of the skin and its derivatives playing a basic role in integrated nervous system physiology, including creativity. And beyond the individual him/herself, there is the niche which he or she occupies, which fosters adaptation to the environment. The construction of the niche and its modifications over time are the principle expression of creativity, at every level. Hope that made sense. John

John, maybe we could assume the same evolutionary perspective. If it is plausible, we could say that the creativity of an animal species is measurable via the new conduct by which it confronts changes in its environment. A possible natural genealogy of creativity can be reconstructed starting from the large differences in behavior between the animal species. Indeed, while some species display more rigid, stereotypical and instinctive behavioural repertoires, others are more plastic and flexible when faced with a novelty and ecological limits. Clearly, in a context in which changes are slow, forms of behaviour which are rigid and specialized (that is to say dependent upon instinctive tactics regulated by genetic memories) are certainly advantageous, but the lack of rapid adaptation, when facing sudden changes, could be fatal to the survival. In contrast, flexible and non-specialized behaviour allows, on the basis of patterns stored, to adopt new models of behaviour while confronting problems which instinctive and stereotyped behaviour can resolve mechanically. 

Mauro, I see your point. Perhaps what you are suggesting could be broken down further to the physiologic response to stress, given that at one end of the spectrum is 'fight or flight' and at the other, the stress hormone epinephrine is needed for learning. So the more reflexive animals are operating under more of the fight or flight mechanism, whereas the more thoughtful types are operating by the learning modality. These are not mutually exclusive, and depending upon the success of their strategy they could move more one direction or the other. Of course all of that should be considered in light of the fact that the effects of chronic stress can lead to depression, both acutely and across generations. John

Dear Mauro I think that is human creativity which has created the neuroscience, neuroimaging and all the theory of brain maps in its image and likeness!

Dear Daniel, I completely agree.

John, I find very striking your vision. On this issue there are many ways distant from each other. Present definitions of creativity refer largely to approaches, abilities and forms of behaviour that generate innovation. Indeed, they correlate creative behaviour to the change in, and maturation of, abilities acquired in the course of evolution. For instance, a child who adapts a toy or other materials and gives them alternative functions is considered creative, as an animal which finds answers to a problem which is not part of its usual stereotyped repertoire. 

Other similar concepts appear to be unsatisfactory. Creativity reflects, either spontaneously or as a result of specific stimuli, the intensifying of perception, of cognition and of other spheres which integrate variables which are not usually associated with one another. In general creativity can be correlated to the capacity of our brain to elaborate a plurality of mental schemes and visions of the world. Every mental function - from perception to memory, from imagination to the representation of reality - is potentially plastic and creative. The source of creativity is to be found in the discrepancy between the world and its representation. Indeed facts and experiences are never represented with exact precision as they are interpreted according to a theory or a vision of reality. If our brain limited its activity merely to the registration of information or to the formation of ‘neutral’ memories – like a computer which categorizes in accordance with logical- mathematical procedures – there would not be sufficient space for creative, divergent, plastic mental processes. For example, the reconstruction of memories, their reinforcement,their incoherence caused by subsequent experiences, their reorganization (both conscious and unconscious), is generated by plastic processes which lead to representations of reality, which on each occasion are different from the initial one.

As regards neural correlations, the classical approach analyses creativity beginning with the various functions of the cerebral hemispheres: that is to say logical-symbolic-linguistic functions which depend largely on the left hemisphere as well as holistic-general-emotional functions which depend generally on the right hemisphere. Naturally, the functions of the hemispheres are even more ample and still little known. Nevertheless, there is no doubt regarding the preponderant role played by the left hemisphere in the symbolic-linguistic activity and in computational activities, to such an extent as to indicate symmetry between biological intelligence and artificial intelligence. 

In my opinion the creativity is ability to make something new, do things in new ways, see things in new ways, express your thoughts, feelings, imagination in new ways (i.e. literature, humor, visual arts, music, dance, sports, engineering, architecture, medicine).

Many brain regions are stimulated by novelty because it is important for our survival. Some by sensory novelty, some by informational novelty. Novelty is considered valuable also by reward coding brain region mOFC, because it might lead us towards potentially useful, richer, more variable (i.e. in nutritious content, genetic variance) resources and flexible adaptations, thus to higher chances to survive in changing environments.

As we are driven by rewards, values, meanings, we are also driven by novelty. All of them are valuable to humans and all induce dopamine signaling that bias us to learn about them, choose them, engage with them, move for them. Creativity, playfulness and curiosity are brain's tools to learn new ways, to get flexible and adaptable to life's unforseen circumstances, to solve problems.

Lateral frontal poles are stimulated by informational novelty, they provoke curiosity (motivation/drive to seek answers to your  questions - via inducing dopamine release in VS). Their role is to question, seek, predict, guess (tool for making models, hypotheses) and find out what is going on around us = the cause and consequences, IF-THEN rules of behavior, contingencies and meanings.

Creativity also leads to finding out new things and new ways and underneath it is human joy of doing something interesting, free, what you or others will enjoy. It is part of value-based choice behavior, and involves joy from achieving goals and dreams (so creativity is also part of goal-directed behavior).

So creative behavior will activate prefrontal cortex, head of caudate (learning the right ways of doing things to reach current goal or solve problem), D1 loop of ventral striatum because of motivation/drive/seeking involved, and also dopaminergic signaling because it confers value and meanings and promotes learning about meaningful things and interrelations.

a reference for lateral frontal pole role in finding out new things and interrelations is in https://www.scienceopen.com/document/vid/13e62385-1fb0-4a49-9a5d-e435baa5b95e?1

Mauro and Karin, I appreciate your perspectives on creativity, but with all due respect, there are much deeper evolutionary origins to creativity, if only we are open to them. In a recent publication (Torday JS, Miller WB. Phenotype as Agent for Epigenetic Inheritance. Biology (Basel). 2016 Jul 8;5(3)) I demonstrated that if epigenetics (acquisition of information from the environment) is combined with Niche Construction (organisms forming their own microenvironments), that the very concept of phenotype as agent emerges. In that context, creativity would be a natural adaptive characteristic. And that concatenation of evolutionary traits can be traced all the way back to the first instantation of life itself, and to the connection between the inanimate and animate realms. Seen from that arc, creativity is a fundamental characteristic of life. To me, that perspective is much more robust than merely describing creative activities and the underlying neurobiology. I would appreciate your thoughts. Best, John

Studies using fMRI, as has shown Alberto Oliverio, have suggested that creativity is correlated to a stand by of the left hemisphere and to an intense activity of associations, allegory and analogies in the right hemisphere, which generate different perspectives. The task of the right hemisphere in the search for new explanations is highlighted by its involvement in a series of functions like perception, musical creation, images and artistic visual creations. The right hemisphere is also involved in the production of associations when using verbal stimuli. It is also known that, because of the chiasmatic structure of the optic passages, perceptive stimuli (like the written word) sent to the right-side visual field are elaborated by the left hemisphere. Furthermore, verbal stimuli elaborated by the right hemisphere are more numerous in analogies and metaphors when compared to stimuli elaborated by the left side of the brain. The immediate resolution of a problem principally involves the right hemisphere. For example, the right answer in a test corresponds to the rapid commissioning of the temporal lobe of the right hemisphere. The commissioning of the right side of the anterior superior temporal gyrus, in particular, is proceeded by a sudden changes in bioelectric activities of the prefrontal cortex, which is implicated involved in several neurophysiological functions. These different competences have led to the attribution of the creative, affective and instinctive functions to the right hemisphere, while the left hemisphere deals with rational thought.

From an evolutionary point of view, the creativity of an animal species is measurable via the new conduct by which it confronts changes in its environment. A possible natural genealogy of creativity can be reconstructed starting from the large differences in behavior between the animal species. Indeed, while some species display more rigid, stereotypical and instinctive behavioural repertoires, others are more plastic and flexible when faced with a novelty and ecological limits. Clearly, in a context in which changes are slow, forms of behaviour which are rigid and specialized (that is to say dependent upon instinctive tactics regulated by genetic memories) are certainly advantageous, but the lack of rapid adaptation, when facing sudden changes, could be fatal to the survival. In contrast, flexible and non-specialized behaviour allows, on the basis of patterns stored, to adopt new models of behaviour while confronting problems which instinctive and stereotyped behaviour can resolve mechanically.

For a capsule theoretical account of brain mechanisms that can enable human creativity, see the section on creativity in "Evolution's Gift: Subjectivity and the Phenomenal World" on my RG page.

Yes, I think it is interesting to add that from an evolutionistic perspective, the central nervous system has selected two neural mechanisms, emotional and cognitive, which process distinct types of information. Once they reach the thalamus, these passages trigger different sequences: an initial one, with emotional significance, is elaborated in the amygdala; the following, with an affective significance, is elaborated in the gyrus cingulate of the limbic cortex and in the ventromedial prefrontal cortex. The cognitive organization, by contrast, contains other limbic structures, largely the hippocampus and the parietal, occipital and temporal lobes of the cortex. Among scholars there is widely agreement on the fact that this system is responsible for long term memory. The functionality of this system depends on the simultaneity of both types of elaboration on the dorsolateral prefrontal cortex. This region takes part in the control of executive functions as well as integrating the information which allows for cognitive superior functions. Furthermore, this brain zone planning strategies for appropriate behavior and execution through the motor cortex and is responsible of the temporal processes, working memory and vigilance that makes possible all higher mental functions.

Creativity is to the regions of the brain as the music is to the keys of a piano.

Again, with all due respect, by trying to understand the principles of consciousness from the existing phenotypic characteristics is, by definition, reasoning after the fact. I know that the conventional approach to biology is descriptive, as it has been since the time of Linnaeus, but if we are to make progress we must become more mechanistic, based on evolutionary principles of pre-adaptational exaptations, i.e. that existing traits evolved from pre-existing cellular-molecular mechanisms. In the case of the brain, and I assume consciousness, it is now known that the link between the invertebrate and vertebrate CNS is the skin, or skin-brain. That homology then leads all the way back to the cell membrane of unicellular organisms. It is that kind of vertical integration that, in my opinion, will yield the answer to the nature of consciousness.

Dear John, I think you mean the homology between Homeobox genes. The same homeotic genes make it possible the development of Arthropods carapace and of the Rodent's rhombencephalon. Nevertheless the answer to the nature of consciousness needs the phenomenological analysis. The knowledge of brain only It is not sufficient to explain the consciousness  and its origin. It is interesting what Plotinus said:it is not the soul that is in the body, but the body is in the soul. Anyway Neurophenomenology could be a way to understand better both consciousness and brain.

Dear Daniele, it's more than just homeobox genes......many genes can be shown to be homologous across species developmentally and phylogenetically. Please see Torday JS. What We Talk About When We Talk About Evolution. Cell Commun Insights. 2015;7:1-15. John

http://pijamasurf.com/2011/10/la-neurobiologia-de-la-narrativa-o-como-contar-historias-es-crear-realidades/

One of the essential characteristics of creativity is the recombination of contents and information available into new associations. As clarified Alberto Oliverio in several papers, creative ideas are the result of previously untried relations between known objects and events mistakenly considered unbeknown to one another. It should be noted, however that it is not deductive reasoning which leads to its creation but rather mental images. Analogies, in particular, reveal similarities and connections between objects and events, which help to bridge the cognitive gap or to resolve problems through the comparison with the repertoire of knowledge previously gained. The theoretical constructs within analogies are similarly found in the outside world which are not representable due to their distance from our senses. This strategy allows many artistic personalities to express concepts which are untranslatable in formal language. This objectification of ideas, which is beyond our perceptions, demonstrates how our mind does not reflect the reality, but builds a new one. Analogies, thus, produces new knowledge from experiences as well as past recollections. Furthermore, it increases the probability of new creations, as opposed to those severe logical results which can inhibit the free associations which flow when our imagination is unbridled. Two areas of the cortex are implicate in the generation of analogies: the prefrontal and the associative areas. It is well known that the associative cortex reformulates in different sequences the elements of an experience lived. For example, knowing someone entails having memorized their face, their voice, the context in which we met them, as well as any emotional reactions we may have had upon meeting the person such as fondness, indifference, aversion and so forth. For instance, beginning with a single signal, such as the tone of voice, the associative cortex reactivates the critical aspects of the face, the emotions involved and so on. In contrast the prefrontal cortex acts as a store room for representations, from which useful elements may be retrieved so as to respond to specific requests. When analogies come into play, the prefrontal cortex selects the information, while the associative areas connects the elements in common.

Analogies are not based upon coded logical inferences, but rather upon logic that, in spite of a number of restrictions, allows us to discover new realities. These restrictions are inevitable to formulate new analogical moulds applicable to inedited phenomena, on the basis of what is noted. In summary, the anthropomorphic monkeys explore reality through a mix of desire to know, investigation and forms of conjectural behaviour which are practical and operational: in much the same way as a child explores all the possible solutions of a game with blocks so as to build castles. The outcomes of the child’s game are most of the times unexpected and hard to distinguish from the assumptions about the effects of his behaviour.

The complexities of creativity reflected in both Leda and Mauro's replies are the reason why I think that creativity has to be addressed at a fundamental level, not by deconvoluting from its overt present form. Smell is one of the most evocative stimuli for memory, for example, and people who cannot form facial gesticulations lose their ability to express their emotions. Conversely, people who have lost limbs experience phantom sensations like pain. In my mind's eye I can only imagine such complex, integrated properties of mind at the level of a unicellular organism, in which stimuli generate holistic responses. Starting from that premise, working our way forward phylogenetically might provide the answer as to what creativity is.

A few years ago, Csikszentmihalyi proposed the concept of “flow”, a state of mind characterized by a sentiment of excited attention and of complete involvement, often erroneously assimilated to creativity. In reality, compared to the difficulty of measuring creativity and its elusive nature, flow is susceptible to experimental analysis both on a quantitative level and as regards the neural correlations. Although the concept of flow has been widely used in a number of contexts, it is characterized by a state of consciousness which is almost automatic, effortless and highly focused. Furthermore, it is often associated with the almost euphoric state in which

operations are carried out without effort or fatigue, to the best of the person’s capacity. “Flow” can concern every mental or physical activity characterized by concentration and commitment, clear objectives, immediate feedback, in proportion to the person’s ability. 

Dear Roberta, thank you for 'connecting the dots' with Csikszentmihalyi's concept of flow as creativity. In my opinion it is consistent with what I have been attempting to express from a cellular-molecular perspective. Briefly, the first instantation of life was the formaton of a cell membrane that separated internal from external, conferring Free Will on the first protocell, but at the same time determining the First Principles of Physiology, contingent on such properties as chemiosmosis (bioenergy), negative entropy (negentorpy) and homeostasis. It is those properties that enable life, but at the same time constrain it......evolution acting to maintain the equipoise controlled by homeostasis; when the organism finds itself in an environment it can no longer tolerate it changes accordingly until it reaches an adaptive equipoise. That process is the basis for creativity, vertically integrated from the local to the CNS......or what Csikszentmihalyi refers to as 'flow'. When the egg is penetrated by the sperm at the time of fertilization, there is a calcium wave that sets embryogenesis in motion. That calcium wave continues throughout life, until the organism dies.......I have attached some of my publications fyi. John

Very interesting, John. Anyway my perspective is a little different. 

The flow may also be considered a temporal interval during which a routine ability or a cognitive function takes place without the interference of the explicit system: a state of lesser activity of the prefrontal lobe that would temporarily “turn off” the analytical capacities of the explicit system.

As is evident in the martial and dynamic arts, the reduction of the time lag between action and reaction corresponds to the reduction of the cognition in a motor programme, to the extent that it becomes simultaneous in exceptional cases. The ‘unaware structures’ take the place of the ego by making use of the abilities acquired with a conscious effort and by favouring the passage from a state of voluntariness to free instinctivity. 

Dear Roberta, I respect your expectation that if you look at the processes of higher neuronal function the principles will emerge. However, I am skeptical for both a priori and a posteriori reasons. A priori, there are First Principles of Physiology that encompass all of physiology, including consciousness. And a posteriori reasoning is reasoning after the fact, by definition. I start from the premise that there is a continuum of consciousness from inanimate to animate, and from the simplest biologic systems to the most complex. If that is correct, then by studying the way that unicellular organisms are aware of their environment, their mind and body being one by definition, that the principles of consciousness can be reassembled, all the way to hominins. I know that this is not the traditional way in which we biologists are trained or funded, but i think that a holistic, synthetic approach needs to be considered, in my opinion......your response is welcomed. John

To restart our conversation I would like to report that recent studies into the function of the striatum have demonstrated how the nucleus accumbens is fundamental in the positive and negative reinforcing of behaviour. Reinforcement, however, can also be interiorized as happen for adults and parents towards the children conducts. Furthermore, the nucleus accumbens operates intervenes in the forms of procedural, declarative and semantics memory. In that sense, this nucleus is an important junction in the network of emotional elaboration of information constructed by the amygdala, the mid-brain nuclei involved in reinforced behaviour, and by cognitive structures such as the hippocampus and the prefrontal cortex. This complex network, which includes other cortical and subcortical structures, highlights its centrality in the elaboration and in the conversion of this information into adequate behavioural answers which could possibly be strengthened. Apart from this, the striatum preempts the gratification of the choices made and indicates the negative effects of those behaviors expected in reward.

Dear Mauro, with all due respect, the analytic approach you are taking is 'reasoning after the fact', whereas by looking in the forward direction developmentally and phylogenetically, the true neurobiological correlates of creativity can be determined. I start with the premise that creativity is an output of the nervous system that starts with unicellular organisms, their cell membrane integrating sensation and response holistically. The homologs of that process can be traced through development with the backdrop of phylogeny. The big breakthrough in the phylogeny of the central nervous system of vertebrates was the article by Nick Holland on 'skin brain', pointing out that the CNS of worms resides in their skin (Holland ND. Early central nervous system evolution: an era of skin brains? Nat Rev Neurosci. 2003 Aug;4(8):617-27.). The molecular homologies between the skin and brain can be used to 'see' how sensation is integrated in the CNS, for example. And that approach can be broadened by seeing the skin as a barrier, like the lung, expanding the candidates for functional molecular homologies. I know that this is not the conventional way of thinking about neurobiology, but I think in this era of genomics that we must broaden our horizons......trying to fit genes into biology out of context is like square pegs in round holes. John

Dear John, I agree with you though our discussion concerns different logical (and physiological) levels. I agree on a broad sense of creativity. In an evolutionary perspective, the creativity of an animal species is measurable via the new conduct by which it confronts changes in its environment. A possible natural genealogy of creativity can be reconstructed starting from the large differences in behavior between the animal species. Indeed, while some species display more rigid, stereotypical and instinctive behavioural repertoires, others are more plastic and flexible when faced with a novelty and ecological limits.

Clearly, in a context in which changes are slow, forms of behaviour which are rigid and specialized (that is to say dependent upon instinctive tactics regulated by genetic memories) are certainly advantageous, but the lack of rapid adaptation, when facing sudden changes, could be fatal to the survival. In contrast, flexible and non-specialized behaviour allows, on the basis of patterns stored, to adopt new models of behaviour while confronting problems which instinctive and stereotyped behaviour can resolve mechanically. So just how does a species to be original and innovative?

Dear Mauro, my point was that there must be fundamental commonalities across species that account for creativity. Seen from the cellular-molecular level, there are specific mechanisms that transcend phenotypes. Or do you question that? I understand that there is a systematic problem we have in biology and medicine, specializing in our own areas of interest, but if we are to make progress I think we need to find common ground through universal cellular-molecular mechanisms that account for simple and complex physiologic traits alike. There is a consensus that multicellular organisms evolved from unicellular organisms, so why not think in terms of a continuum from simple to complex too?

John, there are no relevant differences between us. There is a continuum from simple to complex. For my background I am inclined to give more prominence to the functioning of the central nervous system. For example, think about the relationship between analogies and creative thinking. One of the essential characteristics of creativity is the recombination of contents and information available into new associations. Creative ideas are the result of previously untried relations between known objects and events mistakenly considered unbeknown to one another. It should be noted, however that it is not deductive reasoning which leads to its creation but rather mental images. Analogies, in particular, reveal similarities and connections between objects and events, which help to bridge the cognitive gap or to resolve problems through the comparison with the repertoire of knowledge previously gained. The theoretical constructs within analogies are similarly found in the outside world which are not representable due to their distance from our senses. This strategy allows many artistic personalities to express concepts which are untranslatable in formal language. This objectification of ideas, which is beyond our perceptions, demonstrates how our mind does not reflect the reality, but builds a new one. 

Analogies, thus, produces new knowledge from experiences as well as past recollections. Furthermore, it increases the probability of new creations, as opposed to those severe logical results which can inhibit the free associations which flow when our imagination is unbridled. Two areas of the cortex are implicate in the generation of analogies: the prefrontal and the associative areas. It is well known that the associative cortex reformulates in different sequences the elements of an experience lived.

Researchers have observed that stimuli reported in new or unexpected environments sets in motion the striatum and that the basal ganglia verify the trustworthiness of the forecasts made in the prefrontal cortex. The spheres can be cognitive or motor and it is likely that the basal ganglia are heavily involved in the activation of chemical signals in response to the stress aroused by the dissonance and discrepancy between perceptions and expectations. The striatum, if necessary, also intervenes in the adjustment of any cognitive strategies corresponding to problems of context. In this way the reinforcement mechanisms underlying the cortex, through their interaction with the frontal cortex and the limbic system and striatum play a fundamental role in the programming and choice of adequate actions.

Dear Mauro, with all due respect, I think there is a fundamental difference between what you and I are saying about creativity. You are approaching it in the conventional way of considering the parts and the whole, whereas I am saying that the origins and mechanisms for the evolution of creativity from unicellular to multicellular must be considered in order to understand the ontology and epistemology of the process.

John, that's okay. I do not pretend a consensus between us. 

John and Mauro,

There is an analogy between your debate and the Gabriel Tarde versus Emile Durkheim on the scene of French sociology in the 1880s and 1890s.  Tarde considered that the understanding of the social had to be founded on the understanding of the individuals while Durkheim argued for the understanding of the social based on macro social entities.  

'' And hence, I believe that the only means of explaining this exuberant growth of individual differences upon the surface of phenomena is by assuming that they spring from a motley array of elements, each possessing its own individual characteristics.''

Thanks, Louis. Let a hundred flowers bloom :)

If you will permit me to try a different tack on the subject of 'creativity'. When the sperm fertilizes the egg it generates a calcium burst.......that flow of calcium ions never stops, until the organism dies. Calcium flux is indicative of all life, particularly during neuronal transmission. Creativity can be viewed as the organized stimulation of calcium flux. Understanding how such events occur will provide the insight needed to understand the fundamental nature of creativity.