Escolar Documentos
Profissional Documentos
Cultura Documentos
Abstract
While several mental functions are characterized by parallel computation performed by moduli in the cortex, consciousness is
sustained by a serial global integration: a single scene at a time takes place. Studies on complex systems show that macroscopic
variables, integrating many components activities, undergo fluctuations with an intermittent serial structure when the system is
in a state called “criticality”, characterized by avalanches with inverse-power-law (scale-free) distribution densities of sizes and
inter-event times. Criticality has been established in human brain dynamics during wakefulness. Here we review how the criti-
cal hypothesis is able to explain many recent studies on brain complex dynamics. We focus, in particular, on the global, serial,
intermittent behavior that can be assessed via high-density electroencephalograms, studying transitions between metastable states.
Established as it is during wakefulness, it remained unsolved whether this global intermittent dynamics correlates with conscious-
ness or with a non-task-driven default mode, also present in non-conscious states, like deep (NREM) sleep. Here we show that
in NREM sleep seriality breaks down, and re-establishes during REM sleep (dreams), with unaltered spacial structure, in terms of
complex branching of avalanches. We conjecture that this connectivity is exploited in NREM sleep by neural-bistability waves,
resetting and “parallelizing” portions of the cortex.
Keywords: Consciousness, Sleep, Criticality, Scale-free networks, Intermittency
(d) Small
Serial intermittency Complex networks Worlds in Entropy reduction
the brain
Features of consciousness
(e) Global Workspace explained by the hypothesis (g) high Φ
(f) Operational Modules that awake resting-state brain (integrated information)
works at a critical point
Figure 1: Conceptual map of the literature analysis that represents the minimal background on which the present work is based. Dark boxes with white letters
represent concepts, while arrowed gray boxes with black letters represent results well established in scientific literature of different fields (with different readerships).
Examples of (a) are the works of Diakonos and Contoyiannis and of Turalska et al. [29, 30]; for (b) we make reference to the recent works of Chialvo, for instance
[8]; (c) is a classic result of the theory of critical phenomena, see e.g. the book of Stanley [54]; there is a vast literature about result (d), see e.g. the work of Sporn
[6, 7]; for (e) see the reviews of Baars and Edelman et al. [47]; for (f) we refer to the recent review of Fingelkurts et al. [50]. Finally, literature (g) refers to the
Tononi’s Integrated Information Theory [4, 5]
is robust with respect to a variation of event definition, and also changes value only in correspondence of event occurrences. In
in the presence of superimposed noise [59]. Figs. 2 and 3 a sketch of the two signals ξ(t) is reported. The
We are here interested on global events, i.e., on the “simul- times t0 , t1 , t2 , ... correspond to the occurrence of the events
taneous” occurrences of RTP in different EEG channels. For 0, 1, 2, ..., while τ1 , τ2 , ... are the WTs, i.e., the time interval
each EEG recording, the sequence of coincidences, or (concur- between the events 0 and 1, the events 1 and 2 and so on. In
rent) Multi-Channel RTPs (MC-RTPs), is obtained from single- particular, we have:
channel RTPs via the introduction of two thresholds: The first
(a) Asymmetric Jump (AJ) rule:
one, ∆tc , defines the maximum time distance for two single-
the walker makes a positive jump (ξ(tn ) = 1) in correspon-
channel RTPs (from different channels) to be considered con-
dence of each event n, otherwise it stands (ξ(t) = 0). Then,
current; the second one, Nt , defines the minimum number of
ξ(t) is a sequence of pulses of constant intensity.
concurrent single-channel RTPs required for a MC-RTPs to be
recorded as a global event. Since events that have a distance less (b) Symmetric Jump (SJ) rule:
than ∆tc are considered to be simultaneous, ∆tc must be small. as in the AJ rule, but the walker can make positive or nega-
We herein use ∆tc = 4 ms, equal to the instrumental sampling tive jumps in correspondence of an event: ξ(tn ) = ±1. The
time, and Nt = 5. Nt < 5 suffered from first-neighbor effects, sign ± is chosen with a coin tossing prescription.
and Nt > 5 resulted behavior similar to that reported, but with
decreased statistics. +1 +1
0.5
Detrended Fluctuation Analysis. The diffusion scaling H of
the two random walks introduced above is estimated by means
SJ of Detrended Fluctuation Analysis (DFA) [62]. We briefly re-
call the main steps of this method:
0
1 2 3 µ • For a discrete time L = 4, 5, ..., the time series of the dif-
fusion process X(t) is split into not-overlapping time win-
Figure 4: Diffusion scaling H vs. complexity index µ for SJ and AJ walking dows of length L: [kL + 1, kL + L]. The window number is
rules: AJ (continuous line), SJ (dotted-dashed line).
given by [M/L], i.e., the integer part of M/L, being M the
total length of the time series.
Then, from the artificial signal ξ(t) the diffusion variable of the
• For each time window [kL+1, kL+ L] (k = 0, 1, ..., [M/L]),
CTRW is defined as follows:
the local trend is evaluated with a least-squares straight
j=t
X line fit: X k,L (t) = ak,L t + bk,L ; kL < t ≤ (k + 1)L.
X(t) = X0 + ξ( j) ∆t , (1)
j=0 • The fluctuation is derived in the usual way: X ek,L (t) = X(t)−
X k,L (t) = X(t) − ak,L t − bk,L ; kL < t ≤ (k + 1)L.
being ∆t the sampling time of the experimental time series.
The scaling properties of these random walks were exten- • For a given time scale L, the mean-square deviation of the
sively investigated in several papers (see [38, 36, 37] for a brief fluctuation is calculated over every window:
review) by applying the analytical methods of CTRW. Here we
are interested in the scaling exponent H of the second moment (k+1)L (k+1)L
1 X e2 1 X 2
F 2 (k, L) = Xk,L (t) = X(t) − X k,L (t) (5)
2 L t=kL+1 L t=kL+1
σ2 (t) = h X(t) − X i ∼ t2H , (2)
where X is the mean value of X(t). • Finally, an average over the windows is performed:
Analytical expressions of the scaling H as a function of the [M/L]
complexity index µ were determined in the case of renewal WTs 1 X
F (L) =
2
F 2 (k, L) (6)
with inverse power-law tail: ψ(τ) ∼ 1/τµ . These expressions [M/L] k=0
H = H(µ) are reported in Fig. 4 and summarized in the follow-
ing: H
In the case of a self-similar process, it results: F(L) ∼ L . Then,
µ/2 ; 1<µ<2 by defining z = log(F(L)) and y = log(L), it is possible to apply
a least-squares straight line fit:
HAJ = 2 − µ/2 ; 2≤µ<3
(AJ) (3)
z = Hy + C ,
(7)
µ≥3
1/2 ;
where C is a constant.
(µ − 1) /2 ; 1<µ<2
HS J =
(SJ) (4) Improvement of statistical accuracy in DFA. Given a time
1/2 ; µ≥2
series of total length L, the DFA evaluation is reliable up to
Both rules give a normal scaling H = 1/2 for µ ≥ 3, corre- about L/10 and this is due to the lack of statistics in the long-
sponding to normal (Gaussian) diffusion. For the SJ rule this is time regime. However, we do not have only one time series, but
true also in the range 2 < µ ≤ 3, while AJ rule is super-diffusive several independent time segments, each one separated from the
(H > 1/2) in all the interval 1 < µ < 3. On the contrary, the SJ others by at least one artifact or phase shift in the original EEG
rule is sub-diffusive (H < 1/2) for 1 < µ < 2. We note that, if recording. Several DFA curves can be obtained, one for each
the WTs comes from a Poisson process, the value of H is again time segment, and then averaged to get a mean DFA curve. In
1/2 and, in the long-time, we have a Gaussian diffusion. this way, we are able to compute DFA up to a time given by the
6
maximum among the values Li /10, that is maxi (Li /10), where i a time from an underlying set of parallel scenes, and only
runs over all time segments and Li is the total duration time of this selected scene comes into consciousness;
the i-th time segment. Actually, the statistical accuracy remains
stable up to a time given by mini (Li /10) and then decreases for 4. In conscious states, there’s a competition among cooper-
longer time scales. In fact, the number of segments entering ative global integration and autonomous fragmentation;
the average decreases very rapidly when approaching the time the interplay of these two components constitutes the
scale maxi (Li /10). A common method to improve the statistics metastable regime of brain dynamics and determines the
is glueing different segments together, in order to have a reliable complex intermittent behavior in the EEG field [50, 64, 5].
P
DFA curve up to a time ( i Li )/10. This was done for instance 5. the renewal fractal process derived from EEG data, which
for coding segments in DNA [62]. is defined by the sequence of renewal RTP events, is a par-
We improved the statistical accuracy on longer time scales, ticular serial process, as only a global metastable state (gi-
without the risk of making the running window explore spu- ant cluster) at a time takes place and the short-time RTP
rious signal, i.e. that belong to different segments. Firstly, events mark the death of a metastable state and the birth of
for each sleep phase, we evaluated the minimal duration time: a new one [38, 23].
Lm = mini (Li ); then, for each segment, we computed the DFA
up to time Lm ; finally, we performed the average over all the From the above observations, we are then lead to make the fol-
segments. Note that Lm is not only 10 times greater than lowing assumption:
mini (Li /10), but it is also greater than maxi (Li /10). With this The renewal point process describing fractal intermit-
approach, a much better accuracy on long time scales is ob- tency, which is experimentally defined in EEG data by the
tained. In fact, even if the statistical accuracy is low for the sequence of global RTP events with inverse power-law dis-
segments with the shortest duration times, the number of seg- tributed WTs, is a correlate of consciousness.
ments entering the average is greatly increased in the time range We validate this assumption by comparing different states of
between mini (Li /10) and Lm = mini (Li ), as all segments always consciousness in healthy subjects during sleep. In Section 6
enter in the average operation. we have already given a description of the dataset and of the
methods used to analyze the EEG data, which can be summa-
rized as follows: (a) segmentation and artifact removal; (b) RTP
7. Results and discussion
detection, global brain events (c) computation of event-driven
Criticality has been found both in neuronal networks (models random walks (SJ and AJ) and estimation of second moment
and in vitro, see Refs. [22, 24]) and human brain [8] by inves- scaling H by applying DFA.
tigating the spatial and structural complexity, while temporal The diffusion scaling H of the SJ rule is definitively H =
complexity, i.e., time intermittency, in brain EEG was investi- 0.5 for all time segments and subjects (data not shown for the
gated in our previous papers (see Refs. [38, 23]). In particular, sake of conciseness). This is a signature that the complexity
from the analysis of EEG data in resting state (wakefulness) index µ is greater than 2. In Fig. 5 we show the square root
condition we found that the brain (RTP) events introduced by of the second moment σ(t) for the AJ rule, averaged over all
the authors of Refs. [51, 52] are driven by an underlying re- subjects and nights and over the time segments of sleep cycle I
newal fractal point process with well-defined scaling proper- as explained at the end of Section 6.
ties (fractal intermittency). As already said, it is not clear The second moment scaling H switches from an anomalous
if fractal intermittency is uniquely associated with conscious- diffusion scaling (H = 0.75) in the case of (pre-sleep) wake
ness or with a non-task-driven default mode activity [57], also and REM phases to a normal diffusion scaling (H = 0.5) in
present in non-conscious states like deep (NREM) sleep. To deep (SWS) sleep. Inverting Eq. 3, this means that in wake
clarify this point, let us summarize some observations about and REM phases, which are conscious states, we get an aver-
consciousness: age value µ = 2.5, thus giving fractal intermittency and long-
range correlations, whereas in the deep (SWS) sleep phase we
1. the conscious brain is associated with an emerging “gi- get µ > 3. We recall that normal diffusion (H = 0.5) is also
ant cluster” or Global Workspace [47]) that co-exist with in agreement with a Poisson condition, i.e., with exponentially
clusters of any size having scale-free size distribution, in distributed WTs or, more realistically, with an exponential cut-
analogy with what happens in critical systems (see e.g. off emerging at relatively short WTs and, thus, with short-time
[22, 24] for dynamical avalanches, or [63] for oscillator correlations.
synchronization); The normal diffusion regime during SWS phase could be ex-
2. conscious scenes are unitary and occur serially: only one plained in terms of the fragmentation of the Global Workspace
scene at a time takes place [47]; into local, independent, functional units working in parallel,
which is a condition known to be associated with the lack of
3. consciousness is a sequence of metastable states (giant consciousness. Notice that the fragmentation is related to the
clusters), which reflect rapidly adaptive selection mech- large number of Sleep Slow Oscillations (SSOs) during SWS
anisms in perception and memory; in the consciousness [65, 66], which determine a reset of the neuronal activity by
theory of Baars [47], the Global Workspace is an emerging means of a hyper-polarizing wave putting most neurons in a
serial process that, in some way, selects only one scene at down-state, i.e., a state far from the activation threshold of the
7
100 100
10 P(N)
σ(t)
10-1
1
0.1 10-2
10
0.01
10
0 2
10
3
10 10
4 5
10 10-3
t (ms) REM
SWS
10-4 WAKE
WAKE N2
REM i=1.9
1 SWS 10-5
H=0.75
H=0.5 10-6
1 10 N 100
3 4 5
10 10 t (ms) 10
Figure 6: Probability density function P(N) of the number of concurrent events
Figure 5: Asymptotic time range in the DFA computed for AJ rule applied to for different stages, limited to the first sleep cycle. Concurrency is defined in
different sleep phases (cycle I). Continuous and dashed lines are a guide to the the text, and holds within a tolerance time ∆t, i.e. adopting the prescriptions
eye for the slopes H = 0.75 and H = 0.5, respectively. In the inset we report of previous works [22, 38, 23]. Here ∆t = 4.0 ms. WAKE refers to pre-sleep
the entire time range over which the DFA has been computed. Notice that, in wakefulness. REM, N2 and SWS correspond to the different sleep stages as
the short-time range, the DFA of the three phases (WAKE, REM and SWS) are described in the section ”Methods”. Dashed line is a guide to the eye corre-
essentially superposed, all displaying normal diffusion. sponding to an inverse-power law with index ν = 1.9.
membrane potential. This is also called “electrical silence”. Fi- of N electrodes as follows
nally, from a purely descriptive point of view, we can conclude X X
that the result of Fig. 5 demonstrates that the scaling H, and the CN ≡ ξt(i) ξt( j) · · · ξt(k) , (9)
t i< j···<k
associated complexity index µ, could be proposed as a reliable
indicator of conscious states. The interpretation of these results where we consider all N-uples {i, j, · · · , k} of electrodes, so that
deserve further investigations that, however, will be the focus
of future research work. CN
P(N) = P . (10)
Surprisingly, the distribution of avalanche sizes does not N CN
change when comparing wakefulness, NREM and REM sleep. The result of P(N) illustrated in Fig. 6 thus suggests that the
This means that, at variance with the intermittent behavior, wakefulness functional connectivity is preserved both during
the global topological features of the brain neuronal network NREM and REM sleep. We can rigorously state that even in
are independent from the particular brain condition (wakeful- deep sleep spontaneous patterns emerge, with the same size dis-
ness, REM, NREM). Fig. 6 illustrates the distribution P(N) of tribution of those emerging during wakefulness or REM sleep.
avalanche sizes N, namely the number of electrodes simultane-
ously undergoing an RTP, within time tolerance ∆t = 4.0ms.
The distribution, confirming previous results [23], decays as 8. Heuristic explanation based on physiology
1/N ν with ν . 2. No difference in the distribution is visible
as a function of sleep phases. In [67] it was shown how neural functional connectivity de-
Let us see heuristically how distribution P(N) is related to the creases in deep sleep, compared to wakefulness, after sublim-
topology of functional connectivity. We recall that this prop- inal Trans-cranial Magnetic Stimulation (TMS). It was there-
erty is classically derived from cross-correlation functions. Let fore hypothesized that a pruned or simplified functional topol-
us for simplicity omit the role of ∆t, and consider concurrences ogy was responsible for the lack of consciousness, as it makes
as if the events in different electrodes were exactly at the same the system drifts away from the critical integration/segregation
time. Since our signal is given by the dichotomous time series balance towards a more segregated state. Along this line, [71]
ξ(t) corresponding to the AJ rule, we only have a nonzero con- recently reported that during NREM sleep the dynamics over
tribution for pairwise correlation functions only when we have different brain areas show an higher degree of independency
a concurrence of events among electrodes. Defining C2 as the (parallel computation) with respect to wakefulness and REM
sum of the integrals of all pairwise correlation functions sleep. An even more recent paper confirms this behavior in
XX anesthesia, i.e. during propofol-induced loss of consciousness
C2 ≡ ξt(i) ξt( j) , (8) [72]. Combining these important experimental results with the
t i< j increasing evidence of criticality in brain functioning (at least
during wakefulness) may lead to think that during NREM sleep
where ξt(i) is the dichotomous signal stemming from channel i the system drifts away from the critical regime. Does lack of
at time t, we can generalize this correlation measure for clusters consciousness mean that the system become supercritical or
8
Event: integra,ng excita,ons Downwards causa,on or SSO-‐driven reset (long arrow)
Figure 7: Schematic description of circular causation with and without consciousness. Left panel describes a classic scheme of circular causation in critical systems.
Downwards arrows denote the dynamics of order parameters or, equivalently, the dynamic emergence of organized activity at a given level of description, which
generates an organized structure at a larger-scale level. Upwards arrows denote the constraints imposed by the emergence of structures within a level to a smaller-
scale level. The figure is inspired by Fig.1 of [50], where the smallest scale level is the microscopic one, either associated to neural small structures or to qualia (the
atoms of cognition); the second level is mesoscopic, associated to larger neural formations or to cognitive objects; the third is associated to a macroscopic neural
level, or to phenomenal scenes. Here a further level is introduced, making it explicit the fact that, at criticality, activity clusters continuously form (events) and
break, with only one giant cluster present at the time. Conversely, it also explicits the unitary nature of consciousness, and the emergence of a global workspace. In
the right panel we see the modifications induced by the possibility of neural bistability, which is normally induced by large fluctuations: Till the level of phenomenal
scenes the circular causation described by criticality is conserved; when a giant cluster is being formed the consequential large fluctuations directly reset the system
at the microscopic neural level (long downwards arrow). As a consequence the global workspace never survives in its unitarity. This is graphically underlined by
the use of a dashed line for the largest-scale.
subcritical? The question, at the light of our results, is a lit- coupling among neurons. As a result, it is also possible that the
tle more complicated and further aspects have to be taken into system stays globally critical, drifting from a critical condition
proper consideration. The explanation we propose, illustrated to another. As a matter of fact, an indication of global critical-
in Fig. 7 aims at conciling the results presented herein and the ity is given by the distribution P(N) of quake sizes that remains
known literature, also taking into account that in biology a new unaltered throughout all sleep phases.
form of criticality may take place, i.e. the extented criticality The stability of P(N) with respect to sleep phases may be at
advocated by Bailly and Longo [73]. According to this view, odds with the results of [67]. It has to be said, however, that we
some form of criticality always takes place in biological sys- are herein looking at spontaneous dynamics involving networks
tems, that continuously drift from a critical point to another (for also composed of thalamic neurons, rather than stimulated in a
a review see [74]). top-down fashion, hence shunting the thalamus, via TMS as in
The onset of sleep is characterized by a reduction of neuro- [67].
transmitters and hence of neural activity in the reticular ascend- Sleeping brain is characterized by cellular bi-stability. Neu-
ing system, connecting the brain stem to the cortex. This is fol- rons during sleep are able, due to the opening of calcium-
lowed by a sensory deafferentation. Both mechanisms reduce dependent potassium channels, to hyper-polarize down to nega-
the high frequency activity in the cell yielding a reduction in the tive potentials as low as −80mV. This state, is normally referred
amount of random activity onto synapses, or, in other words, to as “down state” or “electrical silence”. The reason of this be-
of fluctuations. We expect therefore that the system becomes havior is under debate. Some explanations are given by differ-
super-critical, i.e. more synchronized. On the other hand, the ent amounts of chemicals at the end of a prolonged wakefulness
reduction in neurotransmitters may also decrease the effective (s-process) [75]. For our purposes it is sufficient to recognize
9
that after the onset of sleep certain neurons live in a thermody- be devoted to the study of pathological forms of unconscious-
namic state such that an afferent activity may locally generate ness, as it is not clear whether the route to unconsciousness due
the down state. When a coordinated pool of nearby neurons to sleep may be a good model for understanding these condi-
globally undergo the electrical silence, a giant EEG event takes tions. It is foreseeable that epileptic-related unconsciousness
place, called Sleep Slow Oscillation (SSO) [65, 66] which is may share some mechanisms with sleep, while for other con-
able to diffuse to a large portion of the scalp. SSO events mark ditions like traumatic brain injuries (with wide-spread axonal
the death of metastable states, and can be safely assumed to be injury) this similarity is expected to break down.
renewal, since they reset the memory in vast areas of the scalp.
Moreover, they are normally triggered by high levels of neural [1] Y. Nir, G. Tononi, Dreaming and the brain: from phenomenology to neu-
rophysiology, Trends Cogn. Sci. 14 (2010) 88-100.
activity, normally detected as events by our procedure. [2] M. Gell-Mann, Consciousness, Reduction, and Emergence, Ann. New
At the REM sleep onset, the abrupt and intense release of York Acad. Sci. 929 (2001) 41-49; T.E. Feinberg, Neuroontology, neuro-
acetilcholine at the cortical level resets consciousness by block- biological naturalism, and consciousness: A challenge to scientific reduc-
ing potassium channels [68] hence preventing neural bistabil- tion and a solution, Phys. Life Rev. 9 (2011) 13-34.
[3] G. Werner, From brain states to mental phenomena via phase space tran-
ity. On the other hand, the cholinergic activation of cortical sitions and renormalization group transformation: proposal of a theory,
and sub-cortical circuits yields a metabolic mapping typical of Cognitive Neurodynamics 6 (2012) 199-202.
sensory/motor-emotional interaction. Indeed, emotional struc- [4] D. Balduzzi, G. Tononi, Integrated Information in Discrete Dynamical
Systems: Motivation and Theoretical Framework, PLoS Comput. Biol. 4
tures, such as amygdaloid complex, parahippocampal cortex, (2008) e1000091.
anterior cingulate, and sensory/motor structures such as pri- [5] G. Tononi, Consciousness as Integrated Information: a Provisional Man-
mary motor cortex, somatosensory cortex, visual cortex, and ifesto, Biol. Bull. 215 (2008) 216-242.
auditory cortex are overactivated with respect to NREM sleep [6] E. Bullmore, O. Sporns, Complex brain networks: graph theoretical anal-
ysis of structural and functional systems, Nat. Rev. Neurosci. 10 (2009)
and, although non consistently reported, to wakefulness [69]. 186-198; E. Bullmore, O. Sporns The economy of brain network organi-
Finally, Massimini et al. [70] demonstrated that during REM zation, Nat. Rev. Neurosci. 13 (2012) 336-349.
sleep the cortical response to TMS stimulation spreads over the [7] O. Sporns, G. Tononi, G.M. Edelman, Theoretical Neuroanatomy: Re-
cortex as in wakefulness. lating Anatomical and Functional Connectivity in Graphs and Cortical
Connection Matrices. CerebralCortex 10 (February 2000) 127141; for a
Putting everything together, we infer that after sleep onset, popular review see C. Zimmer, 100 Trillion Connections, Scientific Amer-
the main difference with the critical dynamics of wakefulness ican 304 (2011) 58-63 (2011) and references therein.
and REM stands at the highest level of dynamical integration [8] D.R. Chialvo, Emergent complex neural dynamics, Nat. Phys. 6 (2010)
among different areas (Figure 7). At that level, in the waking 744-750 (2010); D. Fraiman, P. Balenzuela, J. Foss, D.R. Chialvo, Ising-
like dynamics in large-scale functional brain networks, Phys. Rev. E 79
brain the dynamics is globally integrated, namely a whole sys- (2009) 061922.
tem dynamics exist that, via downwards causation, is in turn [9] M.G. Kitzbichler, M.L. Smith, S.R. Christensen, E. Bullmore, Broad-
shared by all areas (left panel of Figure 7). After the sleeping band Criticality of Human Brain Network Synchronization, PLoS Com-
onset, a globally emerging large fluctuation actually puts differ- put. Biol. 5 (2009) e1000314.
[10] N.W. Hollingshad, M. Turalska, P. Allegrini, B. J. West, P. Grigolini, A
ent areas at rest, while, in between these resetting events, differ- new measure of network efficiency, Physica A 391 (2012) 1894-1899.
ent areas undergo independent dynamics (right panel of Figure [11] M. Turalska, E. Geneston, B. J. West, P. Allegrini, P. Grigolini, Front
7). In other words, the very same (or a similar) functional con- Physio. 3 (2012) 52:1-7.
[12] L. De Arcangelis, C. Perrone-Capano, H.J. Herrmann, Self-organized
nectivity that during wakefulness or REM, as a result of a large criticality model for brain plasticity, Phys. Rev. Lett. 96 (2006)
fluctuation (over a threshold), involves the global workspace, 028107:(1-4).
during SWS becomes a traveling Slow Sleep Oscillation that [13] S. Zapperi, L.K. Baekgaard, H.E. Stanley Self-organized branching pro-
spreads through the cortex [66], and therefore resets the areas cesses: mean-field theory for avalanches, Phys. Rev. Lett. 75 (1995) 4071-
4074.
that initiated the fluctuation as well as other areas. This makes [14] E. Lovecchio, P. Allegrini, E. Geneston, B.J. West, P. Grigolini, From
it impossible for the different areas to establish the long-range Self-Organized to Extended Criticality, Front. Physiol. 3 (2012) 98 doi:
correlations observed in the genuinely critical states when con- 10.3389/fphys.2012.00098.
sciousness takes place. In other words, the highest level of inte- [15] D. Sornette, Critical phenomena in natural sciences, Springer-Verlag,
Berlin, 2006.
gration cannot correspond to a global workspace, and the brain [16] H.J. Jensen, Self-organized criticality, Cambridge University Press, Cam-
activity is fragmented into parallel dynamics in between SSOs. bridge, 1998.
These different mechanisms, however, make use of the same, or [17] P. Bak, C. Tang, K. Wiesenfeld, Self-organized criticality: an explanation
similar, functional connectivity that is a remaining sign of the of the 1/f noise Phys. Rev. Lett. 59 (1987) 381 ; P. Bak, C. Tang and
K. Wiesenfeld, Self-Organized Criticality Phys. Rev. A 38 (1988) 364
global critical dynamics happening during wakefulness. We re- (1988).
mark that our focus on events makes it possible to study how [18] A.M. Turing, Computing machinery and intellingence Mind 59 (1950)
the independence of neural activities in the different areas, al- 433-460.
ready established in literature [71, 72], is able to take place, [19] D.R. Chialvo, P. Bak: Learning from mistakes Neuroscience 90 (1999)
1137-1148.
exploiting, through the bistability wave called SSO, the very [20] G. Werner Viewing brain processes as critical state transitions across level
same connectivity that is able to maintain the sleep, but also to of organization: neural events in cognition and consciousness, and gen-
reinstate the critical conscious state of wakefulness. eral principles Biosystems 96 (2009) 114-119.
[21] E. Tagliazucchi, D.R. Chialvo, The collective brain is critical, arXiv:
We remark again that the scenario stemming from this work 1103.2070v1 [q-bio.NC] (2011).
refers to the only physiological lack of consciousness, namely [22] J.M. Beggs, D. Plenz, Neuronal avalanches in neocortical circuits The
dreamless sleep. Further research and ad hoc experiments will Journal of Neuroscience, 23 (2003) 11167-11177; D. Plenz, T.C. Thia-
10
garjan, The organizing principles of neuronal avalanches: cell assemblies [47] B.J. Baars, A cognitive theory of consciousness, Cambridge University
in the cortex? Trends Neurosci., 30 (2007) 101-110. Press, Cambridge, UK, 1998; G.M. Edelman, J.A. Gally, B.J. Baars, Bi-
[23] P. Allegrini, P. Paradisi, D. Menicucci, A. Gemignani, Fractal complexity ology of Consciousness, Front. Psychol. 2 (2011) 4.
in spontaneous EEG metastable state transitions: new vistas on integrated [48] S. Franklin, A. Graesser, A Software Agent Model of Consciousness,
neural activity, Front. Physio 1 (2010) 128. Consciousness and Cognition 8 (1999) 285301.
[24] G.L. Pellegrini, L. de Arcangelis, H.J. Hermann, C. Perrone-Capano, [49] G.M. Edelman, Neural Darwinism: The Theory of Neuronal Group Se-
Activity-dependent neural network model on scale-free networks Phys. lection. Basic Books, New York, 1987; G.M. Edelman The remembered
Rev. E 76 (2007) 016107. present, Basic Books, New York, 1989: G.M. Edelman, Naturalizing con-
[25] E. Novikov, A, Novikov, D. Shannahoff-Khalsa, B. Schwartz, J. Wright, sciousness: a theoretical framework. Proc. Natl. Acad. Sci. USA 100
Scale-similar activity in the brain, Phys. Rev. E 56 (1997) R2387-R2389. (2003) 5520-5524; G.M. Edelman, G. Tononi A universe of conscious-
[26] P. Grigolini, M.G. Pala, L. Palatella, R. Roncaglia, Towards the thermo- ness : how matter becomes imagination, Basic Books New York, 2000.
dynamics of localization processes, Phys. Rev. E 62 (2000) 34293436. [50] A.A. Fingelkurts, A.A. Fingelkurts, C.F.H. Neves, Natural world physi-
[27] M. Lukovic, P. Grigolini, Power spectra for both interrupted and perennial cal, brain operational, and mind phenomenal spacetime Phys. Life Rev. 7
aging processes, J. Chem. Phys. 129 (2008) 184102. (2010) 195249.
[28] S.B. Lowen, M.C. Teich, Fractal renewal processes generate 1/ f noise, [51] A.A. Fingelkurts, and A.A. Fingelkurts, Operational Architectonics of the
Phys. Rev. E 47 (1993) 992-1001. Human Brain Biopotential Field: Towards Solving the Mind-Brain Prob-
[29] Y.F. Contoyiannis, F.K. Diakonos, Criticality and intermittency in the or- lem, Brain and Mind 2 (2001) 261-296.
der parameter space, Phys. Lett. A 268 (2000) 286; Y.F. Contoyiannis, [52] A.Y. Kaplan, A.A. Fingelkurts, A.A. Fingelkurts, B.S. Borisov, B.S.
F.K. Diakonos, A. Malakis, On the intermittent dynamics of critical fluc- Darkhovsky, Nonstationary nature of the brain activity as revealed by
tuations ,Phys. Rev. Lett. 89 (2002) 035701. EEG/EMG: methodological, practical and conceptual challenges. Signal
[30] M. Turalska, B.J. West, P. Grigolini, Temporal complexity of the order Process. 85 (2005) 2190-2212.
parameter at the phase transition, Phys. Rev. E 83 (2011) 061142. [53] A.A. Fingelkurts, A.A. Fingelkurts, Brain-Mind Operational Architecton-
[31] P. Manneville, Intermittency, self-similarity and 1/f Spectrum in dissipa- ics Imaging: Technical and Methodo- logical Aspects Open Neuroimag.
tive dynamical systems, Journal de Physique (France) 41 (1980) 1235- J. 2 (2008) 73-93.
1243. [54] H. E. Stanley Introduction to phase transitions and critical phenomena,
[32] D. R. Cox, Renewal Theory, Methuen & Co. Ltd., London, 1962. Oxford University Press, London, 1971.
[33] P. Paradisi, P. Allegrini, F. Barbi, S. Bianco, P. Grigolini, Renewal, mod- [55] F. Lombardi, H.J. Herrmann, C. Perrone-Capano, D. Plenz, L. de Ar-
ulation and blinking quantum dots, AIP Conf. Proc. 800 (2005) 92-97. cangelis, Balance between Excitation and Inhibition Controls the Tem-
[34] S. Bianco, P. Grigolini, P. Paradisi, Fluorescence intermittency in blink- poral Organization of Neuronal Avalanches, Phys. Rev. Lett. 108 (2012)
ing quantum dots: renewal of modulation? J. Chem. Phys. 123 (2005) 228703.
174704. [56] E. Tagliazucchi, P. Balenzuela, D. Fraiman, D.R. Chialvo, Criticality in
[35] P. Paradisi, R. Cesari, D. Contini, A. Donateo, L. Palatella, Characterizing large-scale brain fMRI dynamics unveiled by a novel point process anal-
memory in atmospheric time series: an alternative approach based on ysis. Front. Physio. 3 (2012) 15, doi: 10.3389/fphys.2012.00015.
renewal theory, Eur. Phys. J. Special Topics 174 (2009) 207-218. [57] M.E. Raichle, A.M. Mac Leod, A.Z. Snyder, W.J. Powers, D.A. Gusnard,
[36] P. Paradisi, R. Cesari, A. Donateo, D. Contini, P. Allegrini, Scaling laws G.L. Shulman, A default mode of brain function, Proc. Natl. Acad. Sci.
of diffusion and time intermittency generated by coherent structures in USA 98 (2001) 676-682.
atmospheric turbulence Nonlinear Proc. Geoph. 19 (2012) 113-126. [58] P. Allegrini, D. Menicucci, R. Bedini, A. Gemignani, P. Paradisi, Com-
[37] P. Paradisi, R. Cesari, A. Donateo, D. Contini, P. Allegrini, Diffusion plex intermittency blurred by noise: Theory and application to neural dy-
scaling in event-driven random walks: an application to turbulence, Rep. namics, Phys. Rev. E 82 (2010) 015103 (2010).
Math. Phys. 70 (2012) 205-220. [59] P. Allegrini, P. Grigolini, B. J. West, A dynamical approach to DNA se-
[38] P. Allegrini, D. Menicucci, R. Bedini, L. Fronzoni, A. Gemignani, P. quences, Phys. Lett. A 211 (1996) 217.
Grigolini, B.J. West, Spontaneous brain activity as a source of ideal 1/f [60] E.W. Montroll, Random walks on lattices, Proc. Symp. Appl. Math., Am.
noise, Phys. Rev. E 80 (2009) 061914:1-13. Math. Soc. 16 (1964) 193-220.
[39] O.C. Akin, P. Paradisi, P. Grigolini, Periodic trend and fluctuations: The [61] G.H. Weiss, and R.J. Rubin, Random Walks: Theory and Selected Appli-
case of strong correlation Physica A 371 (2006) 157-170. cations, Adv. Chem. Phys. 52 (1983) 363-505.
[40] P. Allegrini, F. Barbi, P. Grigolini, P. Paradisi, Aging and renewal events [62] C.-K. Peng, S.V. Buldyrev, S. Havlin, M. Simons, H.E. Stanley, A. L.
in sporadically modulated systems Chaos, Solitons & Fractals 34 (2007) Goldberger, Mosaic organization of DNA nucleotides, Physical Review E
11-18. 49 (1994) 1685.
[41] S. Bianco, P. Grigolini, and P. Paradisi, A fluctuating environment as [63] S.H. Strogatz, From Kuramoto to Crawford: exploring the onset of syn-
a source of periodic modulation Chem. Phys. Lett. 438 (4-6), 336-340 chronization in populations of coupled oscillators, Physica D: Nonlinear
(2007). Phenomena 143 (2000) 1-20 (2000).
[42] P. Paradisi, R. Cesari, P. Grigolini, Superstatistics and renewal critical [64] G. Tononi, G.M. Edelman, Consciousness and complexity Science 282
events, Cent. Eur. J. Phys. 7 (2009) 421-431. (1998) 1846-51.
[43] O.C. Akin, P. Paradisi, and P. Grigolini, Perturbation-induced emergence [65] M. Massimini, R. Huber, F. Ferrarelli, S. Hill, G. Tononi, The Sleep Slow
of Poisson-like behavior in non-Poisson systems, J. Stat. Mech.: Theory Oscillation as a Traveling Wave, J. Neurosci. 24 (2004) 6862-6870; A.
Exp., (2009) P01013, doi: 10.1088/1742-5468/2009/01/P01013 . Piarulli, D. Menicucci, A. Gemignani, U. Olcese, P. d’Ascanio, A. Pin-
[44] P. Allegrini, M. Bologna, P. Grigolini, B.J. West, Fluctuation-dissipation gitore, R. Bedini, A. Landi, Likeness-Based Detection of Sleep Slow
theorem for event-dominated processes, Phys. Rev. Lett. 99 (2007), Oscillations in Normal and Altered Sleep Conditions: Application on
010603; G. Aquino, M. Bologna, P. Grigolini, B. J. West, Beyond the Low-Density EEG Recordings IEEE Trans. Biomed. Eng. 57 (2009) 363-
Death of Linear Response: 1/f Optimal Information Transport, Phys. 72; D. Menicucci, A. Piarulli, U. Debarnot, P. d’Ascanio, A. Landi, A.
Rev. Lett. 105 (2010) 040601:1-4; G. Aquino, M. Bologna, B. J. West, Gemignani, Functional structure of spontaneous sleep slow oscillation ac-
P. Grigolini, Transmission of information between complex systems: 1/f tivity in humans, PLoS one 4 (2009) e7601 (2009).
resonance, Phys. Rev. E 83 (2011) 051130:1-12. [66] D. Menicucci, A. Piarulli, P. Allegrini, M. Laurino, F. Mastorci, L. Se-
[45] P. Allegrini, P. Paradisi, D. Menicucci, R. Bedini, A. Gemignani, L. Fron- bastiani, R. Bedini, A. Gemignani, Fragments of wake-like activity frame
zoni, Noisy cooperative intermittent processes: From blinking quantum down-states of sleep slow oscillations in humans: New vistas for study-
dots to human consciousness, J. Phys.: Conf. Series 306 (2011) 012027. ing homeostatic processes during sleep, Int. J. Psychophysiol. (in press)
[46] L. Silvestri, L. Fronzoni, P. Grigolini, P. Allegrini, Event-Driven Power- doi:10.1016/j.ijpsycho.2013.01.014.
Law Relaxation in Weak Turbulence, Phys. Rev. Lett. 102 (2009) 014502 [67] M. Massimini, F. Ferrarelli, R. Huber, S.K. Esser, H. Singh, G. Tononi,
(2009); P. Allegrini, M. Bologna, L. Fronzoni, P. Grigolini, L. Silvestri, Breakdown of Cortical Effective Connectivity During Sleep. Science
Experimental Quenching of Harmonic Stimuli: Universality of Linear 309(5744) (2005) 2228-2232.
Response Theory, Phys. Rev. Lett. 103(2009) 030602. [68] M. Rudolph, J.G. Pelletier, D. Paré, A. Destexhe, Characterization of
11
synaptic conductances and integrative properties during electrically in-
duced EEG-activated states in neocortical neurons in vivo, J. Neurophys-
iol. 94 (2005) 2805-2821.
[69] J.A. Hobson, REM sleep and dreming: towards a theory of protocon-
sciousness, Nat. Rev. Neurosci. 10 (2009) 803-813; J.A. Hobson, E.F.
Pace-Schott, The cognitive neuroscience of sleep: neuronal systems, con-
sciousness and learning, Nat. Rev. Neurosci. 3 (2002) 679-693.
[70] M. Massimini, F. Ferrarelli, M. Murphy, R. Huber, B. Riedner, S.
Casarotto, G. Tononi, Cortical reactivity and effective connectivity dur-
ing REM sleep in humans, Cogn. Neurosci. 1 (2010) 176-183.
[71] M. Boly, V. Perlbarg, G. Marrelec, M. Schabus, S. Laureys, J. Doyon,
M. Pélégrini-Issac, P. Maquet, H. Benali, Hierarchical clustering of brain
activity during human nonrapid eye movement sleep, Proc. Natl. Acad.
Sci. 109 (2012) 5856-5861.
[72] L.D. Lewis, V.S. Weiner, E.A. Mukamel, J.A. Donoghue, E.N. Eskandar,
J.R. Madsen, W.S. Anderson, L.R. Hochberg, S.S. Cash, E.N. Brown,
P.L. Purdon, Rapid fragmentation of neuronal networks at the onset
of propofol-induced unconsciousness, Proc. Natl. Acad. Sci. USA 109
(2012) E3377-E3386.
[73] F. Bailly, G. Longo, Mathematics and the Natural Sciences. The physi-
cal singularity of Life, College Press/World Scientific, Singapore, 2011
(preliminary version, in French: Hermann, Paris, 2006).
[74] G. Longo, M. Montévil, The Inert vs. the Living State of Matter: Ex-
tended Criticality, Time Geometry, Anti-Entropy - An Overview, Front
Physio. 3 (2012) 39:1-8.
[75] A.A. Borbély, A two process model of sleep regulation, Human Neurobi-
ology, 1 (1982) 195-204.
12