The Ancestry of Reason – updates

        Research moves on all the time, and new findings confirm and expand on old ones, or sometimes modify or contradict them.  Here I aim to provide a guide to some of the most interesting new discoveries and ideas relevant to consciousness and its evolution, and to any particularly interesting new books on the subject.   There will also be the occasional thought that has occurred to me since the book went to press.


Ann. Rev. Psychol.    Annual Review of Psychology

Curr. Biol.   Current Biology

Nat. Neurosci.   Nature Neuroscience

Nat. Rev. Neurosci.   Nature Reviews: Neuroscience

PNAS   Proceedings of the National Academy of Sciences

TICS   Trends in Cognitive Sciences

TINS   Trends in Neurosciences

#  Indicates latest additions


The study of consciousness has reached a level which offers plenty to write about and new books on the subject appear regularly.   From Michael Gazzaniga, the eminent neuroscientist particularly associated with the study of split brains and divided consciousness comes The Consciousness Instinct: Unravelling the mystery of how the brain makes the mind   Farrar, Strauss & Giroux 2018.   The philosopher Daniel Dennett, author of several  books on consciousness, has recorded his latest thoughts in From Bacteria to Bach and Back  W. W. Norton & Co. 2017.  Especially interesting are his ideas about Richard Dawkin’s ‘memes’ – of which languages, Dennett points out, are outstanding examples.   And since I particularly enjoyed Stanislas Dehaene’s Consciousness and the Brain: Deciphering How the Brain Codes Our Thoughts (Viking Press 2014) I shall include it here, although it dates back to a little before The Ancestry appeared.  

It’s known that the report from a sensory area of neocortex must activate prefrontal cortex if it is to achieve consciousness.  However, similar levels of input don’t consistently produce the same result.  After studying the success or decay of weak messages a team in Stanley Dehaene’s laboratory confirmed that the variation is attributable to variation in the pre-stimulus state of the brain.  B. Vugt et al.  The threshold for conscious report: Signal loss and response bias in visual and frontal cortex  Science Vol.360. p.537  4 May 2019

# Two new books about consciousness have been published this autumn.  I find both somewhat disappointing in that they focus on philosophical theories of consciousness, rather than on the neuronal machinery which the eminent neuroscientists who wrote them are so well qualified to describe.  And curiously, neither author mentions that evidence that a conscious sensory experience only happens when the  messsage reaching prefrontal triggers a returning message to the pathways which delivered it, thus setting up that all-important maintained circulation of impulses which weaves the stream of sensory inputs into a whole.  (The return message is more likely if there is also input to prefrontal from the alerting systems of the older, lower brain, if the incoming data has matched up to a significance-bearing schema in what I have called the schema-storage areas of the temporal cortex, or if the new information is relevant to whatever the prefrontal cortex is already dealing with.)

In The Feeling of Life Itself: Why consciousness is widespread but can’t be computed  (MIT Press 2019) Christof Koch argues that Integrated Information Theory is the most useful model around.  The essence of this, if I understand it correctly, is that large brains bring huge amounts of information together through extensisve interconnections among circuits, and consciousness is the result.  Digital computers, on the other hand, are ‘built out of circuit components with sparse connectivity and little overlap among their inputs and their outputs’  and don’t need consciousness to unite their operations – hence the book’s subtitle.  But the crucial point, surely, is that brains evolved to promote the survival of their owners and of the genes that created them, and consquently give a lot of space to inputs from the internal and contact senses.  The memory that is consciousness allows these inputs to be related to those from the outside world even if they don’t arrive simultaneously or operate on the same timescale.  Machines only need to deal with a few basic internal situtations – and it’s surely best they stay that way.

Joseph LeDoux’s The Deep History of Ourselves: The four-billion-year story of how we got conscious brains (Viking 2019) provides, in the first half, a succinct account of the probable origins of life and the developments that led from there to vertebrates and vertebrate brains.  Consciousness only arrives in the second half.  Some of the account of how consciousness happens seems to agree with that in The Ancestry, but it’s difficult to be sure because it’s largely phrased in terms of ‘higher order states’ and ‘lower order states’, rather than clearly defined brain areas.  However, rash though it may be for an unqualified amateur to disagree with a highly regarded professor, I suspect that my ideas about exactly what happens in temporal cortex and what in prefrontal are slightly different from his.

The two authors have strongly opposed views on consciousness in other species.  Koch is confident that other mammals enjoy conscious experience too, since they are closely related in evolutionary terms, have similarly constructed nervous systems, and their behaviour is often remarkably similar to ours.  LeDoux, in contrast, condemns the idea of consciousness in other animals as anthropomorphism and roundly berates those scientists who suggest that it exists.  However a few pages later he says this doesn’t mean that mammals and birds lack minds, ‘if by “mind” we mean the abilitiy to think, plan and remember.’  So perhaps he equates consciousness with consciousness of self, more usually defined as the most advanced form of consciousness, rather than with sensory and emotional consciousness.

I, of course, think that this problem is largely solved by my hypothesis that consciousness first emerged in relation to the business of traveling profitably through complex habitats – in other words the business of building and using a cognitive map.  Later it would expand to promote other forms of activity.  To give two examples, it’s hard to imagine juvenile carnivores practising pouncing on prey if they didn’t have conscious visual experience to tell them what had gone wrong when they were unsuccessful.  And surely it’s conscious visual and aural feedback that allows a mother and mobile infants to maintain proximity to each other while simultaneously engaging in another activity, such as browsing or playing.

Since the whole point of a brain is to promote the actions most likely to lead to the survival of its owner and their genes it’s not surprising if our perceptions are shaped by the potentials for action the current environment offers.  Ingeniously recording brain activity while their subjects perceived and navigated virtual architectural scenes, some promising an easy or at least a possible way through, some obviously impassable, Djebbara et al. found that even the early activity in sensory brain areas varied according to the potential for movement.   Z. Djebbara et al.  Sensorimotor brain dynamics reflect architectural affordances  PNAS Vol. 116 p.14769  16 July 2019

The clues to where consciousness is at work

# I wrote in the book that sensory consciousness emerged first in relation to mapping the environment and finding efficient routes through it, and later evolved to direct other activities.  Activities such as motor play and the maternal care of mobile young, I suggested, reveal more advanced forms of sensory consciousness.  In fact,  I think I should have said, these such behaviours could only evolve where the basic machinery of consciousness already existed.


In broad terms there are three different classes of synapse (junction) between neurons.  The transmitter that is released may be excitatory – increasing the chances of the next cell firing – or inhibitory, reducing them.  Or it may be something more properly called a neuromodulator, which modifies the effect of transmitters.  There are quite a few types of transmitter, however, and many types of neuromodulator.  There are also several different types of receptor for each of them, so the possible effects are numerous.

The larger and more complicated the brain the longer the chains of excitatory and inhibitory neurons.   An excitatory cell may excite an inhibitory one, causing it to inhibit another inhibitory one, thereby relelasing the next neuron in line from inhibition ……  clearly the possibilities are enormous, and they have been increasingly exploited as brains grew larger.

In complex brains especially, a delicate balance is needed between excitation and inhibition.  R. Rubin et al.  Balanced excitation and inhibition are required for high-capacity, noise-robust neuronal selectivity  PNAS Vol. 114 E9366  31 October 2017   But there is a puzzle here, since some of the synaptic connections of cortical neurons, which have been thought to store the results of learning by growing stronger through use, have turned out to be distinctly volatile.  G. Mongillo et al. Intrinsic volatility of synaptic connections – a challenge to the synaptic trace theory of memory  Curr.Op. Neurobiol. Vol. 46 p.7  October 2017   However the same authors have found that, although excitatory synapses significantly outnumber inhibitory ones, the latter play a bigger role in storing information.  As long as the overall balance between excitation and inhibition is not disrupted the network can operate efficiently, despite the  inconsistency of excitatory synapses.  G. Mongillo et al.  Inhibitory activity defines the realm of excitatory plasticity  Nat. Neurosci. Vol. 21 p.1463  October 2018.  #  Further confirmation of the role of inhibitory connections in learning comes from S. Krabbe et al.  Adaptive disinhibitory gating by VIP interneurons permits associative learning  Nat. Neurosci. Vol. 22 p.1834 November 2019.

Inhibitory neurons come in several different varieties, each defined by containing some protein  that the others don’t (such as the Vasoactive Intestinal Peptide-expressing neurons above).  The significance is gradually being unravelled.   M. E. Hern & R. A. Nicoll  Somatostatin and parvalbumin synapses on to hippocampal pyramidal cells are regulated by distinct mechanisms  PNAS Vol. 115 p.589  9 January 2018.   Meanwhile a study of cell types in the visual cortex and part of the motor cortex of mice has shown that the excitatory neurons differ more noticeably between the two areas than the inhibitory ones do.  B. Tasic et al. Shared and distinct transcriptomic cell types across neocortical areas  Nature Vol. 563 p.72  1 November 2018


How learning happens is one of the most interesting questions in neuroscience, and prompts a great deal of research.  Much of it focuses on the hippocampus, the seahorse-shaped structure nestling in what’s known as medial temporal cortex – the bit that curls inward on either side of the brain.  It was here it was discovered that neuronal connections strengthen with use and weaken with idleness, something that also happens in other brain areas, with varying timescales and varying degrees of permanence.  Brief  synaptic potentiation keeps a flow of impulses circulating (if prefrontal cortex has become involved) back around the parts of the sensory brain where it was initiated, allowing a succession of sensory inputs to be preserved for long enough for patterns spread out over time to be perceived.  Most importantly this allows us and other animals to build up a map of the environment and move around it in a purposeful manner.

In building a map through exploration it’s important, of course, to preserve the order in which sensory events occur.  A recent study investigated this aspect of  the hippocampus and closely linked adjacent structures further – N. C. Heyworth & L. R. Squire  The nature of recollection across months and years and after medial temporal lobe damage  PNAS Vol. 116 p.4619  5 March 2019.   Subjects with damage to the medial temporal lobe were taken on a 25 minute guided walk during which eleven planned events occurred, and their recollections of the walk were compared with those of controls immediately afterward, after one month, or up to two and a half years later.  The most outstanding difference was that the memory-impaired patients reported the incidents of the walk in random order, whereas the control group preserved the order.

Another important function to which the hippocampus contributes is recording what is where.  C. E. Connor & J. J. Knierim  Integration of objects and space in perception and memory   Nat. Neurosci. Vol. 20 p.1493 November 2017.  This is one of several articles in an issue focussed on spatial cognition.

Subjective experience suggests that the experiences and discoveries we think about later, or dream about in subsequent sleep, are the things more likely to stick in long-term memory.  Sure enough, many experiments have shown that the exchanges between hippocampus and neocortex which occur during new experience may be repeated during periods of idleness and in sleep, and do indeed serve to fix the new information in long-term memory.   The replays take the form of condensed, high-frequency bouts of neuronal firing termed ripples, which consolidate the connections initiated earlier.  Prolonged ripples create the sturdiest records.   A. Fernandez-Ruiz et al.  Long-duration hippocampal sharp wave ripples improve memory  Science Vol. 364 p.1082  14 June 2019.   Studies in young rats show how these replays of neuronal activity develop as the brain matures.  L. Muessig et al. Co-ordinated emergence of hippocampal replay and theta sequences during post-natal development  Curr. Biol. Vol.29 p.834  4 February 2019.

#  Memory is selective, however, and it turns out that there is an editing mechanism which can suppress hippocampal activity.  S. Izawa et al.  REM sleep-active MCH neurons are involved in forgetting hippocampus-dependent memories  Science Vol. 365 p.1308  2o September 2019

Hippocampal sharp wave ripples, meanwhile, also accompany recollection, occurring just before a memory is successfully accessed.  A. P. Vaz et al.  Coupled ripple oscillations between the medial temporal lobe and neocortex retrieve human memory   Science Vol. 363 p.975  1 March 2019.   Moreover exceptionally detailed recollection of autobiographical events has been found to correlate with exceptionally strong connectivity between prefrontal cortex and hippocampus.  V. Santangelo et al.  Enhanced brain activity associated with memory access in highly superior autobiographical memory PNAS Vol. 115 p.7795  24 July 2018.  

Obviously the first step in creating a coherent idea of surrounding space is to take eye movements into the account, a point considered by M. Nau et al.  How the brain’s navigation system shapes our visual experience  TICS Vol.22 p.810 September 2018   I suspect that damage to the hippocampal formation, leading to difficulty in connecting the views gained from a series of fixations, may account for the difficulty some dementia sufferers have in finding their way around even in homes they have lived in for decades.

The hippocampus allows us to find patterns in inputs spread out over extended periods of time.  The most valuable pattern, for almost any animal, must be the shape of its environment, knowledge which enables it to head as directly as possible for whatever it currently needs.  Another area involved in the mapping process is the adjacent entorhinal cortex, where cells fire in such a way that the animal’s position is registered in terms of a grid covering all the area within sensory reach.  An important bit of data here concerns the locomotion that takes the animal from one junction on the grid to the next.  A contribution to the understanding of this system comes from M. G. Campbell et al.  Principles governing the integration of landmark and self-motion cues in entorhinal cortical codes for navigation  Nat. Neurosci. Vol. 21 p.1096  July 2018.  Another type of entorhinal cell has been found to fire when an object within the surrounding space is at a specific distance and direction.  Object-vector coding in the medial entorhinal cortex  O. A. Hoydal et al.  Nature Vol 568 p.400 18 April 2019

Remarkably, it has been possible to create a computer programme which mastered the art of navigating around a modest area using a grid system based on the entorhinal one.  A. Banino et al.  Vector-based navigation using grid-like representations in artificial agents  Nature  Vol. 557 p.429  17 May 2018

Our view of surrounding space is often limited by buildings, hedges or other barriers to vision and locomotion.  Increasing attention is being given to how the brain records such things.   X. Gofman et al.  Dissociation between postrhinal cortex and downstream parahippocampal regions in the representation of egocentric boundaries  Curr. Biol. Vol. 29 p.2751 5 August 2019;   Q. He & T. I. Brown  Environmental barriers disrupt grid-like representations in humans during navigation  Curr. Biol. Vol. 29 p.2718  5 August 2019  

Alas, in us long-lived humans the area in which hippocampus and entorhinal cortex are found seems to be particularly vulnerable to wearing out.  M. Stangl et al.  Compromised grid-cell-like representations in old age as a key mechanism to explain age-related navigational deficits  Curr. Biol. Vol. 28 p.1108  2 April 2018.  (This too may be relevant in dementia – see earlier.)

Another contributor to navigation and the creation of the cognitive map is retrosplenial cortex, which seems to record the large-scale patterns, including those beyond sensory reach, and the overall geometry of a habitat.   An addition to this picture comes from A. S. Alexander & D.A. Nitz  Spatially periodic activation patterns of retrosplenial cortex encode route sub-spaces and distance travelled   Curr. Biol. Vol. 27 p.1551  4 February 2019

#  And now A. S. Persichetti and D. D. Dilks find that the retrosplenial cortex response to buildings – and no doubt other landmarks – is strongly affected by their position in the landscape.  Distinct representations of spatial and categorical relationships across human scene-selective cortex  PNAS Vol. 116 p.21312  15 October 2019

Mapping the environment must be an ancient form of learning.  There are many other sorts.  A skill that evolved comparatively recently is learning how to use a flexible muscle system to the best advantage.   How it happens is becoming clearer. A. J. Peters et al.  Reorganisation of corticospinal output during motor learning  Nat. Neurosci. Vol. 20 p.113  August 2017   And as might be expected, new patterns of neuronal activity also emerge in the brain.  E. R. Oby et al.  New neural activity patterns emerge with long-term learning  PNAS Vol. 116 p.15210  23 July 2019

Here too sleep serves to consolidate learning, and even a very brief period of rest can help, it turns out.   M. Bonstrup et al.  A rapid form of offline consolidation in skill learning  Curr. Biol. Vol. 19 p.30219  2019

Some forms of learning are so well understood now that scientists have been able to create a response to a previously insignificant odour by co-ordinating stimulation in mouse olfactory cortex with stimulation in two areas concerned with reward and discouragement.   G. Vetere et al.  Memory formation in the absence of experience  Nat. Neurosci. Vol. 22 p.933 May 2019


The primary visual cortex has been most extensively studied in primates and in cats, where neurons favouring lines of a particular orientation cluster together, forming columns.  A new study finds that this holds good in carnivores generally and in ungulates – but not in rats.   M. Weigand et al.  Universal transition from unstructured to structured neural maps  PNAS Vol.114 E4057  16 May 2017   This looks like another example of the general principle that the more complicated brains get the more delicately organised they have to be.

Early studies of visual cortex, confined to recording from only a few neurons, indicated that different aspects of the visual input were processed through separate channels.   Now that much more sophisticated recording equipment is available it seems that a good many neurons in V1 (in macaque monkeys) are sensitive to both colour and orientation.   A. K. Garg et al.  Rethinking primary visual cortex function  Science Vol. 364 p.1275  28 June 2019    More is being uncovered, too, about a later stage of the pathway through visual cortex, V4.   Neurons in the upper layers here extract more detail to pass on to the next processing stage, while neurons in deep layers seem to be more concerned with behavioural relevance and the possible need for attention.  W. W. Pettine et al.  Laminar segregation of sensory coding and behavioral readout in macaque V4  PNAS Vol. 116 p.14749  16 July 2019

A thought about colour vision: the magnificent array of colours we experience, produced by comparing the inputs of just three wavelength-sensitive types of photoreceptors and one less fussy one, is perhaps a particuarly telling example of what the extended processing of sensory input that leads to consciousness can achieve.  Some invertebrates have a much wider range of colour-tuned receptors – dragonflies, for instance, and mantis shrimps.  At least one variety of the latter has twelve sorts.  But the shrimps don’t learn to distinguish among different blends of input.  H. H. Thoen et al.  A different form of color vision in mantis shrimps   Science Vol. 343 p.411  24 January 2014   It seems pretty certain that where there is such a wide range of wavelength-tuned receptors each variety is dedicated to eliciting one particular hard-wired response, comparable to the male stickleback’s automatic display indicating territorial ownership at the sight of any patch of red.  The visual systems of some of those brilliantly coloured little reef fish probably work the same way, with receptors pretty precisely matched to the colours displayed by the species.  Shoaling with fellow members of the species is managed, I would guess, by a hardwired response to the distinctive colour patterns they display – blue and yellow stripes, red and green ones, or whatever.

The received wisdom has long been that cone receptors are tuned to different wavelengths of light while the more sensitive rod receptors respond to a wide range and record only luminance.   Now it has been discovered that some species of deep-sea fish have a whole range of rod receptors tuned to different wavelengths (38 in the most extreme case) – which probably function to detect the bioluminescent signals that are quite common in the deep ocean’s inhabitants.   Z. Musilova et al.  Vision using multiple distinct rod opsins in deep-sea fishes   Science Vol. 364 p.588  10 May 2019

It has been becoming ever clearer that activity in sensory pathways throughout the neocortex can be influenced by factors such as attention snd emotion.  Now it’s been shown, in the mouse, that even neurons in primary visual cortex registering such basic visual elements as vertical or horizontal lines can fire somewhat differently depending on whether the overall pattern of input is new, or constitutes a view the mouse has seen before.   A. B. Saleem et al.  Coherent encoding of subjective spatial position in visual cortex and hippocampus  Nature Vol. 567 p.158  10 September 2018.  This intriguing result was obtained by an ingenious experiment in which the mouse ran on a revolving ball while a changing scene was projected in front of it, allowing tight control of just what the animal sees and where it thinks it’s going.  The results of several such virtual reality experiments have been reviewed by Liam Drew in Nature:  The mouse in the video game  Vol. 567 p.158  4 March 2019.   This experimental method has even been extended  to study how olfaction guides locomotion.  B. A. Radvansky & D. A. Dombeck  An olfactory virtual reality system for mice  Nature Communications Vol.9  article 839  2018.

Tiny, unnoticed eye movements called microsaccades ensure that fine features of the visual scene are at some point perfectly centred in the receptive fields of appropriate detectors in the visual system.   Schelkova and colleagues confirm how helpful this is for distinguishing very small-scale stimulus-patterns.  N. Schelkova et al.  Task-driven visual exploration at the foveal scale  PNAS Vol. 116 p.5811  19 March 2019.

The micromovements of the eye are far too small to be consciously noticed but I nurse a strong suspicion that that they are what make the eyes of mammals and birds seem lively and bright – in contrast to the glassy look of reptilian and fish eyes – and are part of what attracts infant attention to eyes.   But as far as I know nobody has ever looked for microsaccades in birds, let alone in reptiles, and nobody has checked for unconscious effects on attention.

Mirror Neurons

Mirror neurons, identified first in monkeys then in humans, fire both when an action is performed and when the same action is seen being performed, apparently for the same purpose.  They are found in various parts of both motor and parietal cortex.

The evolution of such neurons must underpin the capacity for imitation clearly seen in many mammalian and avian species.  And what massive consequences this talent has had.  An individual who can imitate can learn what is worth doing without taking the risks involved in  experimentation, so knowledge gained by a single adventurous animal can be passed on from generation to generation painlessly.  Moreover these new behaviours can become established in a very short time, in contrast to those which emerge from genetic changes to brains or muscular systems.  (Indeed the influence may work in the other direction – the learnt new behaviours can create a selective pressure which favours modifications to brain or muscle.)

The capacity for imitation is particularly valuable for infants.  In its simplest form it probably involves only the release of an innate action when the infant sees the parent performing it, so that the infant learns to look for food where the parent looks, and what sort of thing prompts flight in the parent.  This form seems to be present in birds.  I have seen a peahen take a few pecks in a likely piece of earth, which prompted her chick to forage there while she moved on a little.  Something similar has been reported in domestic chickens.  I imagine it’s a very similar mechanism which often causes a human infant only a few days old to smile back at someone who smiles at them, and infant monkeys to perform similar imitations.

This form of imitation seems not significantly different from the way little fishes in shoals tend to follow any individual which makes a sudden turn, so that the change of direction rapidly sweeps though even a large group.  Perhaps we should look for pretty ancient origins for the first mirror neurons.  Once they were established, though, they created a situation in which parental care of the young, providing readily available models to imitate, could become increasingly valuable.

The more complex forms of imitation no doubt evolved as muscle systems grew more complex.  And as brains grew larger infants could build on the sensations created by automatic imitations and develop the ideas which support conscious purpose.  Thus humans and no doubt other primate infants gradually develop a conscious sense of how facial expressions coincide with emotion and mood, and how they operate as a means of communication.

The talent for imitation becomes even more useful in social species, and seems to need no further development.  Malaysian sun bears normally lead solitary lives, apart from a period of about three months when a mother is rearing a couple of infants.  However some young bears that were being rehabilitated before return to the forest readily played together in a social fashion, and numerous instances were observed of two sorts of open-mouth expression – something that sounds rather like the chimpanzee play-face – being reciprocated.   D. Taylor et al.  Facial complexixty in sun bears: exact facial mimicry and social sensitivity  Scientific Reports 9  article 4961   2019   This report seems, incidentally, to add to the evidence that many large-brained mammals can switch between social and solitary life styles as the circumstances make appropriate.  (Coyotes provide a particularly notable example.)

As well as mirror neurons for action there are also emotional mirror neurons – empathy neurons – which are active both when emotion is experienced and when someone else is seen experiencing it.  These too must contribute enormously to infant education, demonstrating what is to be feared, what can be approached confidently.   They’ve now been identified in rats as well as primates.  M. Carillo et al.  Emotional mirror neurons in the rat’s anterior cingulate cortex  Curr. Biol. Vol. 19 p.30322  2019   There is evidence, too, of their existence in at least some birds.  J. E. C. Adriaense et al.  Negative emotional contagion and cognitive bias in common ravens (Corus corax)  PNAS Vol. 116 p.11547  4 June 2019

There is evidence, incidentally, that motor imitation is a built-in urge that has to be suppressed.  By adulthood the instinct is normally completely inhibited and must be released from inhibition when wanted, but damage in a certain brain area can affect the ability to control it.  Patients instructed not to copy an experimenter’s actions have difficulty in complying.  It’s obvious that emotional mirror neurons work in the same way – it’s all to easy to catch an emotion or mood, and it takes practice and self-control to avoid infection.

On being social

In social animals both sorts of mirror neuron are obviously also useful in adulthood, serving to spread useful information through the group.   And if the group must deal with a predator or an attack from a rival group their best chance lies in sharing the same emotion and acting in unison – either all standing up to the threat or all running away.  A mixed reaction reduces the chances of survival for everybody.  The most successful group leader, meanwhile, is likely to be the animal which can not only defeat all rivals but can also powerfully activate emotional and mirror neurons in others when needed.  This ability, which also implies a capacity for attracting attention, is surely the basis of that indefinable quality charisma, and the leader who has it generally makes for a more successful group.

Our capacity for living together in large groups, sharing our ideas, combining our efforts, and submitting to some form of central governance, has clearly contributed greatly to our success as a species.  But the very characteristics that made the vast and rapid expansion of our numbers possible carry dangers in the modern world, for what works well in small groups isn’t always beneficial in large societies.  Emotional mirror neuerons at play in a crowd can produce mass hysteria.  Motor mirror neurons can prompt people to do things they wouldn’t if they stopped to think about it.  Charisma doesn’t necessarily coincide with wisdom.  Now that we are organised into very large societies and equipped with the means of destroying ourselves completely (along with many other species) one of the challenges is to keep mirror neurons under control.

# The downsides of our advanced form of sociability are considered in a new book from the biological anthropologist Richard Wrangham, The Goodness Paradox (Pantheon Books 2019).  He develops the idea – first mooted by Darwin – that we humans have acquired many of the traits of domestic species, and have in effect domesticated ourselves.  In particular we have become easier-going, less prone to get into fights with fellow group members, repressing what he calls ‘reactive violence’.  But language, he suggests, made it easier for lower-ranking males to gang up effectively against the alpha male in early, ape-like societies and dispossess or kill him.  It also made larger social groups possible, which thrived best where there was co-operation.  Conformity became highly valued (?the power of both types of mirror neurons grew) and the members of these large groups were prepared to commit cold-blooded, pre-planned violence on non-conformist members – ‘proactive violence’ – which would become more controlled with the advent of police and law courts.  A parallel development led to organised warfare, in place of the quick, unexpected raids on an unprepared enemy that were more likely in smaller groups.  So the goodness paradox is that we have become nicer in one way and nastier in another.

Language has certainly played an important part in these processes – reference to Robin Dunbar’s observations on the importance of gossip in propagating behavioural norms might be appropriate here.  (Grooming, Gossip and the Evolution of Language  Faber 1996).  But even before language some group leaders will have guided group behaviour more intelligently than others, promoting co-operation and group success.  Not all dominant chimps are the violent bullies Wrangham focusses on – so the process of self-domestication may have begun rather earlier than he seems to think.

As for inter-group violence we must face the fact that the ‘survival of the fittest’ – the strongest, the better organised, the better equipped with weapons – will have played a significant part in our evolution.  Evolution which has led us to a place where we can no longer afford the instinct that produces such conflicts.

The knowledge stores

As we learn about the world around us, and about our own capacities, the knowledge is stored in the neocortex, neatly classified into different areas, just as different subjects are stored in different sections of a library.  But the brain also constructs a complex network of connections between the areas, so that the record of a visual stimulus-pattern is linked to records of the sounds connected with it, of any emotions it might evoke, and so forth.  In humans there is also likely to be a link to a word.  Registering a stimulus-pattern in any one area can thus prepare the way for related input elsewhere, or for producing the appropriate word.  And as we learn to recognise particular stimulus-patterns in ever greater detail the neuronal response becomes ever more finely tuned, the pathway of excitation increasingly refined by new inhibitory connections.

Particularly complex stimulus-patterns carrying several forms of significance activate several areas.   Faces are a notable example, analysed separately for identity, expression, orientation, significance – and new face-sensitive areas keep being discovered.  Landi and Freiwald have found (in rhesus monkeys) two in which, as blurred faces gradually beame clearer, the response to familiar ones ramped up quite suddenly, parallelling the way a face can suddenly be recognised.  The response to unfamilar faces grew more steadily, as in other face-sensitive areas.   S. M. Landi & W. A. Freiwald   Two areas for familiar face recognition in the primate brain   Science Vol. 357 p.591  11 August 2017

#  Most investigations of these schema-storage areas have been concerned with visual input, but a new one looked at odours, recording from the fish equivalent of mammalian olfactory cortex.  As zebra fish were conditioned to associate different effects with different odours the odours were ‘systematically remapped along an axis in coding space that represented attractiveness’ – as revealed by the behaviour of each fish.  T. Frank et al.  Associative conditioning remaps odor representations and modifies inhibition in a higher olfactory brain area  Nat. Neurosci. Vol. 22 p.1844 November 2019

J. S. H. Taylor et al. have analysed the areas involved in recognising written words. Mapping visual symbols onto spoken language along the ventral visual system  PNAS Vol. 116 p.17723 3 September 2019

The neocortical areas which store knowledge of such important things as faces, bodies and landmarks are pretty consistently sited.  How does this orderly arrangement come about?  Several explanations have been put forward, not all mutually exclusive.  My own proposition is that these categories of stimulus-pattern are the ones for which the infant has genetically determined responses – imitation for faces, bodies and voices, grabbing for grabbable objects, and so on – and the hardwired action is reported to a predestined part of the neocortex, which is thereby sensitised to the sensory input which evoked the action.  This hypothesis doesn’t rule out other factors which might be at work, such as the suggestion that the reward infants gain from looking at faces and engaging in social interaction with them may play a role.   L. J. Powell et al.  Social origins of cortical face areas  TICS Vol. 22 p.752  1 September 2018

But the importance of action in perception seems to be underlined by an experiment showing that accuracy in making a quick estimate of the number of visual objects in a display is greatly influenced by the number of fixations made, suggesting that “numerical estimation ability is closely tied to the mechanisms that control eye movements and attention.”  S. J. Cheyette & S. T. Piantadosi  A primarily serial, foveal accumulator underlies approximate numerical estimation  PNAS Vol. 116 p.17729  3 September 2019

Certainly the development of these areas is not dependent on the sensory channel through which the input comes.  The language areas of congenitally deaf people who communicate by means of visual sign language are similarly situated to those of people who use spoken language.  In like manner, it’s now been found, the congenitally blind store knowledge of faces, bodies, objects and scenes in much the same way as the sighted, although their knowledge must be  gained entirely through other senses.   J. van den Hurk et al.  Development of visual category selectivity in ventral visual cortex does not require visual experience  PNAS Vol. 114  E4501  30 May 2017   (This fits well with the long-ago discovery that the blind can draw recognisable pictures of things like tables that they know through touch.)

The output from these areas which deal in subjects such as faces is passed on to yet other neocortical areas which record the asociations among inputs from different senses, such as one towards the front of temporal cortex which responds not only to the appearance which defines an individual but also to the name that goes with it, and any other known aspects of the person concerned.  Y. Wang et al.  Dynamic neural architecture for social knowledge retrieval  PNAS Vol. 114 E3305  18 April 2017

Evolutionary history

Interesting ideas and new discoveries about the past continue to emerge.  It seems that the transition from single-celled animals to multicellular ones may have been less complicated and occurred more often than once thought.  E. Pennisi  The power of many  Science Vol. 360 p.1388  29 June 2018

Our knowledge of the weird and wonderful multicellular animals that had evolved by the time of the Cambrian period will be increased by the discovery of an extensive deposit of remarkably well preserved fossils – including, unusually, some of soft-bodied animals – on a river bank in China’s Hubei province.  It will be fascinating to correlate them with the long-known fossil riches of the Burgess Shale in Canada, also from half a billion years ago.  And now this Canadian fossil-bearing shale has been found to extend for kilometres beyond the original site, and more exciting finds are appearing.   J. Sokol   Cracking the Cambrian    Science Vol. 632  23 November 2018

# Other recent fossil discoveries are modifying old ideas about the emergence of mammals.  J. Pickrell  The Making of Mammals  Nature Vol. 574 p.468  24 October 2019

Darwin’s essential insight is simple enough.  Those organisms best fitted to meet the challenges of survival are likely to breed the most prolifically, and with many descendants comes the best chance that some will adapt to new circumstances and giving birth to new species.  But changing environments and competition for food and living space among the evolving species create all sorts of complexity.  Pringle et al. explored one of the outstanding questions in a long-term experiment making use of 12 small Bahamian islands on which the top predator was a semi-terrestial insect-eating lizard.  They introduced either a ground-dwelling lizard capable of eating smaller lizards as well as arthropods, or a tree-dwelling species, or both, while 4 islands were left as controls.   When all three species were present numbers of both smaller species declined, but the experimenters deduced that this was more because of the competition for living space than because of predation by the carniverous lizards.   The original inhabitants learnt to make more use of the trees, forcing the specialists in arboreal living to retreat to higher levels.   R. M. Pringle et al.  Predator-induced collapse of niche structure and species coexistence  Nature Vol. 570 p.58  6 June 2019

An important aspect of the evolution of Homo sapiens has been an increasing ability to get on with each other, leading to the possibility of ever larger social groupings, which permit the exchanges of ideas that can inspire major technological advances.  All sorts of genetic modifications have contributed to this development – a reduction in the tendency to be aggressive to strangers, an increase in facial expressiveness and so on.   Many of the qualities selected for parallel those selected by us in the domestication of other species, prompting the suggestion that we have in effect domesticated ourselves.   An interesting treatment of this subject is B. Hare  Survival of the friendliest:  Homo sapiens evolved via selection for prosociality  Ann. Rev. Psychol.  2017

Motor matters

Why should there be two locomotory centres in the brain?  I hypothesised in the book that they might govern slightly different forms of locomotion, serving different purposes, such as gentle exploration and hasty escape.  New research suggests that I was partially right.  The different forms are not quite what I guessed at – and there are probably more than two such centres.   V. Caggiano et al.  Midbrain circuits that set locomotor speed and gait selection   Nature Vol. 553 p.455  25 January 2018.

Other interesting papers on motor subjects are  M. N. Economo et al.  Distinct descending motor cortex pathways and their role in movement   Nature Vol. 563 p.79  1 November 2018  and  Z. Gao et al.  A cortico-cerebellar loop for motor planning   Nature Vol. 563 p.113  1 November 2018


Modern lifestyles make considerable demands on attentional systems, and the effects are receiving some interest from neuroscientists, as in M. R. Uncapher & A. D. Wagner   Minds and brains of media multitaskers: Current findings and future directions  PNAS Vol. 115 p.9889  2 Occtober 2010    The effect of multimedia use in early years is also increasingly studied, as in D. A. Christakis et al.  How early media exposure may affect cognitive function: A review of results from observations in humans and experiments in mice  PNAS Vol. 115 p.9851  2 October 2018

Attentional priorities differ, it seems, among individuals in a way that probably reflects both nature and nurture.   Viewing a large series of complex scenes some people fixated earlier and longer on faces, others on text, others on food, and so on.   B. de Haas et al.  Individual differences in visual salience vary along semantic dimensions  PNAS Vol. 116 p.11687  11 June 2019


The neuromodulator dopamine is probably one of the most ancient conveyors of messages between neurons – perhaps it even antedates neurons.  I became rather obsessed with it when I first started studying the brain, until I feared I was placing too much emphasis on it, and told myself to be more objective.  But it has turned out to be deeply involved in so many aspects of brain function – movement, attention, learning, reward, addiction – that it seems fair to deduce it has had a particularly vital role in brain evolution.  Here are some of the latest papers.

B. Enghelhard et al.  Specialised coding of sensory, motor and cognitive variables in VTA dopamine neurons  Nature Vol. 570 p.509  27 June 2019;   A. Mohebi et al.  Dissociable dopamine dynamics for learning and motivation  Nature Vol. 570 p.65  6 June 2019;   M. Sheng et al. Emergence of stable striatal D1R and D2R neuronal ensembles with distinct firing sequence during motor learning  PNAS Vol. 116 p.11038  28 May 2019;   L. Ferren et al.  Dopamine modulates the reward experiences elicited by music  PNAS Vol. 166 p.3793  22 January 2019;  C. E. Stelly et al.  Pattern of dopamine signaling during aversive events predicts active avoidance learning  PNAS Vol. 116 p.13641  2 July 2019

Serotonin (also known as 5-HT) is another extremely influential neuromodulator, and must be at least as ancient as dopamine.  But the latter got a headstart with neuroscientists when it transpired that a shortage of it was a major factor in Parkinson’s disease.   A shortage of serotonin would later be linked to depression, which fired up scientific interest, but the complexities here still seem even more challenging than for dopamine.  However a useful review of the current state of knowledge has been published:  B. W. Okaty et al.  Embracing diversity in the 5-HT neuronal system  Nat. Rev. Neursci.  Vol. 20 p.397 April 2019

The thalamus

The thalamus is a central structure in the brain, part of the diencephalon (the second of the major divisions of the brain, which are based on the way the swelling at the front end of the embryonic spinal cord is initially divided).  In primates and many other species the thalamus is thoroughly buried under the vastly expanded telencephalon, and was consequently very difficult to get at with recording needles until extremely fine ones were developed.  For a long time it was known principally for relaying visual, auditory and somatosensory information to the necortex for further analysis, and for its contribution to shutting off access from sensory systems to the neocortex during sleep.  Now a great deal more is being discovered, though the overall picture is still far from clear.  A valuable exposition of the new evidence is L. Acsady  The Thalamic Paradox  Nat. Neurosci.  Vol. 20 p.901  27 June 2017.

Here are some other recent papers:   S. S. Bolkan et al.  Thalamic projections sustain prefrontal activity during working memory performance   Nat. Neurosci. Vol. 20 p.987  July 2017;   M. M. Hakassa & S. Kastner  Thalamic functions in distributed cognitve control   Nat. Neurosci. Vol. 20 p.1669  December 2017;   G. Pergola et al.  The regulatory role of the human mediodorsal thalamus  TICS Vol. 22 p.1011  November 2018.

The mammalian thalamus has many detectable specialised divisions.  Those of such amphibians as have been studied are much simpler.  I’ve suggested that the structure began as a sort of map, in a self-centred reference frame, of the animal’s current situation, one that came into existence at a period when all actions were pretty much hardwired.  Actions would be reported to the map, and in a way that indicated the position of the stimulus that prompted them – the position in relation to the animal that is.   Probably just enough visual information came from the retina to the proto-thalamus to anchor the map to the external world.  (The visual data which guided the action would be dealt with by the visual centre in the lower brain which in mammals has been so greatly overshadowed by the visual cortex.)   The complex functions of the mammalian thalamus could have evolved, I postulate, from this sort of beginning.

Prefrontal cortex and decision-making

What does the brain do when the rules that guide profitable behaviour change?  An answer comes from M. Sarafyazd & M. Jazayeri  Hierarchical reasoning by neural circuits in the frontal cortex  Science Vol. 364 p.6441  17 May 2019

Addicts fail to control their drug use despite knowing the negative consequences.  This reflects an increase in the power of ‘go’ signals from the orbitofrontal cortex to the basal ganglia and a decrease in the power of suppressive signals from other prefrontal areas.   Y. Hu et al.  Compulsive drug use is associated with imbalance of orbitofrontal- and prelimbic-striatal circuits in punishment-resistant individuals  PNAS Vol. 116 p.9066 30 April 2019  The imbalance may be related to a less than normal growth of the myelin that insulates the nerve fibres in the former pathway during adolescence.   A similar deficit in a parallel pathway from lateral and medial prefrontal cortex can be linked to impulsiveness.   G. Ziegler et al.  Compulsivity and impulsivity traits linked to attenuated developmental frontostriatal myelination trajectories  Nat. Neurosci. Vol.. 22 p.992 June 2019

Decision-making involves ‘working memory’ in which prefrontal cortex plays a leading role.  An impressive use of high-resolution functional MRI has distinguished the activity of upper layers of dorsolateral prefrontal cortex from that of lower layers during a working memory task.  #  E. S. Finn et al.  Layer-dependent activity in human prefrontal cortex during working memory  Nat. Neurosci. Vol. 22 p.1687 October 2019

#  Prefrontal cortex must be vital to conscious thought and reasoning.  ‘The stream of thought can flow freely, without much guidance from attention of cognitive control’ observe J. N. Mildner and D. I. Tamir.  They propose Spontaneous thought as an unconstrained memory process  Trends in NeurosciencesVol. 42 p.763 November 2019

The Default Mode Network

Over the past couple of decades cortical areas have been discovered which are quiet when attention is paid to the outside world and active during remembering and thinking.   It’s now become apparent that they add up to “multiple interwoven networks.”  # R. L. Buckner & L. M. DiNicola  The brain’s default network: updated anatomy, physiology and evolving insights  Nat. Rev. Neurosci.  These networks must surely support, in the authors’ words “a class of processing function dependent on internally constructed rather than externally constrained representations.”

However there is intriguing evidence that the default network is not always suppressed when attention is paid to the external world.  It can be active when aesthetic pleasure is derived from viewing paintings, architecture or landscape.  #  E. A. Vessel et al.  The default mode network represents aesthetic appeal that generalizes across visual domains  PNAS Vol. 116 p.19155  17 September 2019

Parts of the default mode network are also involved in mental imagery.  #  J. Pearson  The human imagination: the cognitive neuroscience of visual mental imagery  Nat. Rev. Neurosci.  Vol. 20 p.624 September 2019


# Acquiring language is clearly likely to have a considerable effect on the structure of the developing brain.  Studies of children who were born deaf and had no access to sign language while growing up are revealing patterns of connectivity very different from those with early language exposure, and cognitive differences too.  L. Boroditsky  Language and the brain  Science Vol. 366 p.13  4 October 2019

F. Huettig & M. J. Pickering suggest that people who learn to read fluently thereby learn to process language more quickly, gain an increased awareness of words as linguistic units, and where homophones are spelt differently can distinguish them more readily.   The learning done through reading, moreover, can be done with less distraction.   Literacy advantages beyond reading: Prediction of spoken language  TICS Vol. 23 June 19

Future directions

One of the most exciting aspects of neurobiology is the increasing understanding of mental disorders, and the promise of reducing the impact they have.  In more general biology, meanwhile, a hot topic is the internal microbiome – the vast array of micro-organisms that live within and on us in profitable symbiosis.  Skolnick and Greig have brought the two subjects together, suggesting that some symbiotic microbes, established in us for untold generations but sometimes lost in these days of liberal antibiotic consumption, may be essential for mental health.  They cite two that contribute to the synthesis of neurotransmitters and one that assists in the excretion of mercury and possibly other heavy metals.  These pathways, they say, ‘have the potential to impact systems relevant to a wide range of neurodevelopmental and psychiatric conditions including autism, depression, anxiety and schizophrenia.’   S. D. Skolnick & N. H. Greig  Potential neuropsychiatric consequences of dysbiosis  TINS Vol. 42 p.151 March 2019

A suggestive new idea about schizophrenia is to be found in L. Barkovitch et al.  Disruption of conscious access in schizophrenia  TINS Vol. 21 p878  November 2017.   And it looks as if pyschopathy might be explained at least partially by a deficiency of emotional mirror neurons.   L. A. Drayton et al.  Psychopaths fail to automatically take the perspective of others  PNAS Vol. 115 p.3302  27 March 2018    Nevertheless they are not incapable of considering the welfare of others, it was established.   Hope for the addicted, meanwhile, is held out by M. Diana et al.  Rehabilitating the addicted brain with trancranial magnetic stimulation  Nat. Rev. Neurosci. Vol. 18 p.685  November 2017

Recovery after stroke involves incorporating new neurons in areas adjacent to the damage.  The process is fuelled by corporal and the concomitant brain activity, and it turns out that brain areas compete for the new neurons (which are produced in the subventricular zone, SVZ)   H. Liang et al.  Region-specific and activity-dependent regulation of SVZ neurogenesis and recovery after stroke  PNAS Vol. 116 p.13621  2 July 2019

Neonatal intensive care wards for premature infants tend to hum with the noise of essential machinery, and often music is played to mask the sound and provide more meaningful sensory stimulation.  Lordier et al. find that music helps to establish important brain circuitry in preterm infants.   L. Lordier et al.  Music in premature infants enhances high-level cognitive brain networks  PNAS Vol. 116 p.12103 11 June 2019