JSA $28 08 8.0-595-54201-4 2 THE SOURCE A basic assumption of neuropsychology ~ grounded in the cinco. anatomical method ~ is that if a particular mental function i performed by a particular brain region, then a complete lesion of that region must result in complete loss of that function. As I have shown, when it comes to consciousness, the cortex fails this test, But things are even worse for the cortical theory of consciousness: lesions elsewhere in the brain obliterate it completely — and very small lesions at that. The physiologists Giuseppe Moruzzi and Horace Magoun established over seventy years ago that consciousness in cats is lost following tiny incisions that disconnect the cortex from the ‘reticulate’ (net-like) core of the brainstem.! This core must be eee 525 million years old, because it is shared by all es to humans. Since Moruzzi and Magoun’s » researchers have confirmed in all manner of species cause coma. For example, David brtinsem stroke a eel identified in human patients with i ‘wo cubic-millimetre ‘coma-specific ntine tegmentum (see Figure 1).’ densely knotted core f He explanations. The first is that ths of the brainstem is where consciousness the hiddi a iowa pring of the mind, the source of i OMSthat itis ike thet J8ak Panksepp, The second els” omer cable for a television set: necess* T Mesop. fete ate to Possib 122The Source or sufcient, and hardly enlighsening i 7 What 0 a st nd is how television works, This is th OU Want n wo sis the 0 WN Gappose the second option is true, By ga "*ilstteam yc. : Stimulating the ae, we might expect to switch OMScioUsHess oy or aor brain. ve might attenuate it in ae ways, like a a A best ising the TV screen to fade. We woutdn — ‘eduction thecurrent broadcast on the fly. And yet an electrode im, ; Tewrite reticular brainstem nucleus of a sixty-five. Planted in pem nucle year-old wom, ¢ eset of Parkinsons disease) reliby coke markable response? The patient’s face expressed Profound sadness within five seconds [...] Although still alert, the Patient leaned to the right, started to cry, and verbally communicated feelings of sadness, guilt, uselessness, and hopelessness, such as “t', falling down in my head, I no longer wish to live, to see am. thing, hear anything, feel anything ...’ When asked why she was crying and if she felt pain, she responded: ‘No, I'm fed up with life, ’'ve had enough ... I don’t want to live any more, Pm disgusted with life ... Everything is useless, always feeling worthless, I’m scared in this world.’ When asked why she was sad, she replied: ‘I’m tired. I want to hide in a corner I'm crying over myself, of course ... I'm hopeless, why am | bothering you?” [...] The depression disappeared less than 9 seconds after stimulation was stopped. For the next five minutes the patient was ina slightly hypomanic state, and she laughed and joked with the examiner, playfully pulling his te She recalled the entire episode. Stimulation [at cae brain site, which was the actual target of the electrode] di Not elicit this Psychiatric response.’ ri in previous c ¢ symptoms ; ti ; dno ious history of psychiatric symP “ny kind, , ipplies to chem! ckade of these ms li ical stimulation OF block 123Tue HIDDEN SPRING brainstem nuclei. Most antidepressants — Serotonin f, in core bral ns whose cell bodies are located in 4 Tegion s ~acton neuro, system called the raphe nuclei (seg py! the reticular activat ved there, as we say. Antipsychotics — 4, Serotonin s sO rons sourced in another part of the blockers — act wa the ventral tegmental area (see Figure out activating syste nxiety drugs ~many of which block . The same apples ani which is produced by neuross sca called noradrenaline, which is p pa age or) in the locus coeruleus complex {also Figure 1), yer another Part of the reticular activating system. All these neurons are clumped together in the reticulate core of the brainstem. Psychiatrists wouldn't tinker with this region of the brain if it merely switcho consciousness on and off, If that were all it did, it would interes, only anaesthetists. The second view must therefore be wrong, Functional neuroimaging of the brain in emotional states points to the same conclusion. Positron emission tomography during states of GRIEK SEEKING, RAGE and FEAR, for example, show that the highest metabolic activity occurs in the core brain- stem (and other subcortical regions; see Figure 9) while the cortex shows deactivation. Functional magnetic imaging during orgasm reveals the same: the haemodynamic activity that correlates with this intensely affective state is almost exclusively located in the midbrain ae brain stimulation, pharmacologic conclusion: the a neuroimaging all point to the os Apparently, therefore, : Sore of the brainstem generates # fe ‘ 10 be necessary for dees? Part of the brain that we kno¥ “aul poner ne ns Consciousness as a whole his * felng In the brevious hone another mental function, name Basten whan woe how feelings pena late our ie with feelings eae else it is meant to be doing, ee a Needs) ich come from within us and ae NESS, Bue jg P Peat to be one of the central i 7 "OW looks as though the ncurologi@ 124The Source eee Happy-neutral Fear-neutral Figure 9 Positron emission tomographic images of four emotional states (courtesy of Antonio Damasio). The upgoing arrows indicate regions of increased activation and the downgoing arrows indicate regions of decreased activation. The highlighted area in the ‘Joy’ image appears to show activation of the SEEKING system. sources of affect and of consciousness are, at a minimum, deeply ‘stangled with one another, and they may in fact be the very same ‘achinery. Contrary to the classical empiricist view, according to uhich consciousness flows in through our senses, and contraty to ‘esatement I quoted from Meynert that was based on that view, “eems the brain does ‘radiate its own heat’. ‘athe should we call the basic medium, this mysterious ee sae pet to well up within us? We cannot call it a‘waking °48Zeman did, since that would requite usto describe dream sane of wakefulness, which i absurd. We canal fasingeg 2°el either, as Moruzzi and Magoun did, ; ‘esctibed show thatitentails intensely qualitative feature 125Tue HIDDEN SPRING os try the third term used in the literature: ‘arousa} neutral word for it. Both waking and q ortddoes not preclude quality ash sitively suggests fecling This am, tern So, let’ mea goods sal; an ing involve arou' ; ‘arousal’ po: ‘level’ does. In facts Ip We have talked about it in straightoryarg ardly put what és arousal out iti behavioural terms ~ for example, the distinctions between com, rand fully responsive wakefulness. Is usualy Y the vegetative ireasured by the Glasgow Coma Scale: tests of the patient’s je. opening responses, their verbal responses to questions and thei motor responses tO instructions and to pain. However, it can also be defined physiologically. An EEG produces graphic tracings Of cortical electrical acti. ity. When left to its own devices (1. 1 ar activating system, even when processing Sen") PV the cortex produces the delta wave pattern, a series of high-ampli- every second (i.¢. a frequency ude waves occurring roughly twice of 2H). When stimulated by the reticular activating system in the absence of sensory input, the cortex typically produces the theta hythm (4-7H2)° or alpha thychm (desynchronised waves at frequencies of 8-13Hz; ‘desynchronised” means erratic). Wher actively processing external information, the cortex typically dis plays the beta (desynchronised, very low amplitude waves with frequencies of 14-24Hz) or gamma pattern (low amplitude wavs with very high frequencies of 25-10oHz). Gamma is the rhytha most commonly associated with consciousness. Nowadays, it is possible also to measure physiological using functional neuroimaging, which literally pict brain aciriy by mapping regional patterns of change in merelt ee nee this technique, with reference °° * i arousal, which i . ee row of the figure shows RE M ay) fexneie aac showed, NOt ee upper brainstem. Neu consciousness, and which is sourced PT . Neuroimaging of some of the basic emotion if disconnected from the 126« The Source 04 oe a time (s) o Hew vtvlio [HS Nl hh NAN a $y ae e d the most “r0used (delta) pattern is shown at the top andthe mos (gamma) pattern at the bottom. The i crvity, The least Fame 10 Typical patterns of cortical EEG ati and beta. Patterns, from top to bottom, are thetay alpha al 127Tue HIDDEN SPRING £ consciousness described above (see Figure 9) en hows the same thing: brainstem arousal, nd of omar vortex becomes conscious only t0 the exten aroused by the brainstem. The relationship between the two ig hice archical; cortical consciousness depends upon brainstem Arousal That is why the delta wave pattern shown at the top of F; — which is not associated with conscious behaviour — jg Benerateq by inteinsc cortical activity, and that is why the gamma ry, shown at the bottom of Figure 10 ~ which is strongly Associated with consciousness — can be driven by the reticular activating system alone. That is also why the drop in physiological arousal shown in the top row of Figure 3 coincides with waning of con. sciousness and sleep onset, whereas the increased arousal shown in the bottom row coincides with reappearance of consciousness in dreaming. These facts are not controversial. Let’s peer more deeply into the actual brain mechanisms involved. To do this, Ineed to introduce a basic distinction between two ways in which neurons communicate with one another. This distinction turns out to be important for consciousness. t that it Igure 1p Most people with a casual interest in the brain know that neurons transmit messages along intricate networks, This process is called Synaptic transmission, because it involves Passing messages across Synapses, the structures by which one neuron conveys signals to another (the word ere Greek for {join — ynapse’ derives from the Gree - Synaptic transmissi vo F ansmission uses molecules called 1H transmitters, which are pas either exciti ssed from one neuron to the neh ing on the me ons Post-synaptic neuron or inhibiting it, depe™” excitatory ae westion (glutamate and aspartate 4 iransmil i Fa (GABA) ae mitters and gamma-aminobutyt'¢ 4 flurry of ent If the downstream neuron is excited! thenesy he Tansmitters, it thee TONS in the Networl NSF Molecules quick vg on 0 Passes its own molecules a r; se areatteh If it isn’t, it doesn’t. There® ck into ly degrade or are taken back 128Yy a The Source i ron, to limi ke cule t the duration of their effect ~ 4 prot << transmission is targeted, bi Sync brain function hatiomeee end api 1 oD, which may explain why it has boett of digital gervous SYStEMs including the cortex. But it is not intcinseally fonscious. In other words, this type of neurotransmission ocean, jn the cortex whether it is conscious or not. And it h: pothing to do with arousal. What fewer people know is that synaptic transmission takes glace under the constant influence of a completely different physiological process. This other type of neuronal activity is Hiled post-synaptic modulation. Unlike synaptic transmission itis messy, inescapably chemical and very different from what happens in a typical computer. It arises endogenously from the ticular activating system (and other subcortical structures, and xn from some non-neurological bodily structures); and it has everything to do with arousal. The central players in this process are a class of molecules led neuromodulators. Unlike neurotransmitters, these mol- ecules spread diffusely through the brain — that is, they are ttkased into the general vicinity of whole populations of neurons vather than at individual synapses.” Instead of passing messages dong specific ‘channels’, they wash over swathes of the network, theeby egulating the overall ‘state’ of the cortex. For example, ings is in a different state in the top Peas ‘er a in Figure 3 (slow wave sleep vs REM sleep) an ve emotional states shown in Figure 9 (GRIEF vs SEEKING tion ae FEAR). In each of these states, it processt cool erently. Thus, if someone calls your name when Y =— ae neuromodulatory molecules can act also as itters, on occurs las next to 129ISBN TuE HIDDEN SPRING sence ery differently from how you reace ‘Act when y OU are asleep, You ety awake” Likewise consider your response t0 an ay -— state of SEEKING versus tite the former state YOU might greet the stranger, and even « EARS i, whereas in the latter state you nk ta 7 ght look rae rersation with thes coe hope they dont notice YOU. iy and Nrion between channel” and State 82. we hand for the ewo ways in which neurons communicate wit Ae and ratte of te cores afets he differential sn the message-passing that goes on within ieths of theme estate adjusts how ‘oud the cite’ cham of spake ex Figure 12). That ib why the same sound ie commu ling your name) i spread widely i the cortex ie wvakefulness but sequestered in the auditory cortex during sleep, and why a stranger arouses one brain network during the FEAR n its channels; ina many anner state and another during SEEKING. This is the crux of what we call arousal. Note, however, that odulated both upwards and downwards, cortical arousal can be m‘ as happens to the point of suppressing tran: every night when we go to sleep ( g -P prefer the term ‘modulation’ ove! determines which synaptic impulses will b sirongly, as in the example of your name are asleep versus awake. Synaptic transmission is binary ( synaptic neuromodulation grades the likelihood that coer fire, It shifts the statistical odds that so appen in them. This probabilistic, analogue adjustmen’ mes rhe through receptors located at several place é : : ath of the neuron. Unlike neurotransmitters> neu it lators have relatively sl affects ~"° eet ly slow-acting and long-lasting © oe i ; cause channels that fie chemicals themselves lingers but PE. yet time. If ire more frequently become more likely © - If you boost som il i ¢ part of the network, it will stay smission completely, which is why some physiologiss i ‘grousal’). Arousal therefore e transmitted, and how being called when you but post on/off, yes/no, 1/0) n set ol agivel mething *! of firing 130—— The Source wart nin (ere en aa ny WWE Lara pL TET Wit ou ae nie Pe tim reat mate ue WPM vy Wt VE Hetero g : Te Th : Hite te ta mitt i Hitt wi VL " H Woe tomMh ob mwa ptm vib borin i meu mn tn nou Te \ EE EW EM rE ee nil Piout to menent wt tt o 500 1000 Time (ns) Figure 11 The image above is a plot of the spike trains of twenty .¢ vertical y axis) over a period of 1.5 neurons (represented on th 1 axis) during presentation of seconds (represented on the horizontal 2 visual stimulus. Stimulus presentation occurs at time o on the x axis (represented by the second vertical line). At this point the neurons, which hea baseline firing rate of 6Hz (on average, when there is no stimulus) incase their average firing rate to 30Hz. A spike train isa sequence in which a neuron fires (= spikes) and does not fire (= silences). This canbe represented as a digital sequence of information: ‘1’ fora spike ad's fora silence, For example, an encoded spike train could read as fuutoroy, The frst woos here represent the latency time between lus presentation and the first spike. The important por ma * fring rates are not determined by the stimulus lone they ae ae ee stimulus, the long-term porentiato ition of the neurons (i.e. whether the stimulus is familar °° of ae and their current level of modulation. The ads is veanaptic modulation is the effect of arousal on canes a isu the stimulus depicted here might evoke n0 87008" Me neurons when the arousal is modulated cownwa fro note is 131Tue HIDDEN SPRING | such time as it is down-modulared. This influences i] su i vor! boosted unt! nd itis a big part of how learning works, Arousal neuroplasticity lessons more deeply in the channels of jous our variou : states inser! e for example, YOu are more likely to remembe, A ourbras you are anxiously trying tO find an unfamiliar des. j when journey i the same place by h, are travelling to y habit, nae n when you > tination thal han _ onal odulators come from? They come from al do neu Where do neon the pituitary, adrenal, thyroid and sex bod, incl Dey + rode various hormones) and the hyporhalana (which produces innumerable peptides). But the conceal source of ‘arousal’ from the brain’s point of view is the reticular activating