I promise the next one will be more personal.

Here's my basic summary of mirror neurons so far:

Mirror neurons are different to ordinary neurons in that they activate both when an individual performs an action, and when s/he observes or sometimes imagines an action. These patterns of activation can be related to physical movement, watching another individual be touched, communicative gestures and, perhaps, emotions. One of the most prominent hypothesis as to the purpose of mirror neurons is the direct-matching hypothesis. This is the idea that MNs are used to compare observed (or imagined) behaviours to internal representations of those behaviours. If the comparison is accurate, the individual making the representation is thought to begin to understand both the physical requirements and the purpose of the behaviour to some extent.

The idea that mirror neurons (MNs) are important in understanding and recreating behaviours/expressions led to many researchers trying to link MN activity to empathy. This primarily involved tests on autistic individuals, who tend to have a much lower level of empathy than unafflicated participants. Many valient efforts have been made, but as far as I can tell within the limited time I have spent examining the area, it is still a work in progress. Up to the present the studies have followed the hypothesis: "If MNs are the mechanism for understanding social and motor behaviours, and autistic individuals have problems with this ability, then MNs may be responsible." Unfortunately the results have been very mixed. Most of the findings from self-report surverys follow something called the "Extreme male brain" hypothesis, in which women report more empathy, men report more systemising and autistic individuals follow an extreme version of the male standard (e.g. Cheng et al. 2008). Unfortunately, a significant proportion of the research shows no real association between empathy, systemising and MN activity during tasks that were designed to reveal such a link if it were there. Mirror neurons do seem to provide some comparison between external stimuli and eternal representations, but that function may not be linked to behaviour, temperament or personality in a straight forward manner.

Confused by the scattered results in the subject of empathy, I looked elsewhere to find a focus of research, and was happy once again to stumble upon some research by a terrific researcher and public figure: Vilayanur Ramachandran. His name popped up all over the place when I was studying the stuff of consciousness, and it appeared he had a lot to say on the topic of mirror neurons also. Specifically, he spoke about the apparant barrier between observing someone else experiencing a sensation, and actually experiencing the sensation for oneself. I highly recommend you read this article he wrote to get an idea for his incredibly engaging style and his way of thinking outside the box in a truly awe-inspiring way ( http://edge.org/3rd_culture/rama08/rama08_index.html ). That article is the source of this quote:

"I also suggest that although these [mirror] neurons initially emerged in our ancestors to adopt another's allocentric visual point of view, they evolved further in humans to enable the adoption of another's metaphorical point of view. ("I see it from his point of view" etc.) This, too, might have been a turning point in evolution although how it might have occurred is deeply puzzling.

There are also: "touch mirror neurons" that fire not only when your skin is touched but when you watch someone else touched. This raises an interesting question; how does the neuron know what the stimulus is? Why doesn't the activity of these neurons lead you to literally experience the touch delivered to another person? There are two answers. First the tactile receptors in your skin tell the other touch neurons in the cortex (the non-mirror neurons) that they are not being touched and this null signal selectively vetos some of the outputs of mirror neurons. This would explain why our amputee experienced touch sensations when he watched our student being touched; the amputation had removed the vetoing. It is a sobering thought that the only barrier between you and others is your skin receptors!"

Sobering indeed. Ramachandran brought up the perfectly sensible question of why YOU aren't feeling a baby grasp your thumb at this very moment. Individuals with phantom limbs may actually feel it, because their skin doesn't tell them otherwise.

Avid viewers of the popular American TV drama "House" may have seen an episode in which the main character cured a patient's phantom pain by constructing a mirrored box with holes for each arm. The patient slid his real arm and his 'phantom' arm into the box, and the mirror made it look to him like he now had two arms again. By being asked to clench his fist(s) he felt like he was releasing tension in both arms, and his pain was gone. Remarkably, this isn't fiction. The mirror box is used in real life phantom pain reduction and was invented by Ramachandran and his his wife, Rogers-Ramachandran : http://en.wikipedia.org/wiki/Mirror_box . MN networks still exist even if a limb is put out of action via amputation or numbing. Ramachadran, Mrs Ramachandran and a graduate student called Laura Case performed an experiment (I can't find the reference, but they mention it several times!) in which they numbed participant's arms and then had them observe an experimenter being touched or picking up an ice cube. Without skin sensations, the participants could feel the touch, and even experience the cold chilling their fingers! Similar phenomenon have been reported regularly in studies of mirror box therapy (e.g. Chan et al. 2007).

Another thrilling feature of Mirror neurons is that their responses depend to some extent on life experience. This was first shown in macaques who showed mirror neuron activation in response to an experimenter using tools to play with food, only after they themselves had learned how to use tools in the same way. In humans similar results have been shown for piano playing and classical ballet dancing amongst many other skills. In fact, is has been shown to be possible to train humans to ‘counter mirror’ and display activity in neurons controlling areas of their body that are different from those being used in external stimuli (i.e. being shown an index finger prodding a piece of paper, could provoke mirror responses in neurons which control the pinky finger instead). If mirror neurons change as a result of experience, then it would suggest that comparing activity to measures of empathy would depend heavily on individual participant’s previous experience of empathy. Something that would perhaps be impossible to reliably measure through self reporting. Combining this work with research on touch MNs (see above) suggests that MN networks are malleable and will change in response to external factors. Lessons learned, loss of limb etc. Even the earliest research into macaques found that some MNs would only activate in response to a very specific copy of an executed movement. It is hard to imagine that the MN networks would be designed to account for adult habits of hand movements which are so informed by learned habit.

One variation of Ramachandran's mirror box. The mind can be tricked by such simple apparatus. Don't believe your sense of self is always limited to your body.

Mirror neuron inhibition seems to be a natural feature of the MN system in humans at least. WE may experience MN activity in response to watching a man do a handstand, but we don't immediately jump to copy the action. This inhibition may be weaker in infants (who tend to copy certain mouth movements despite not even being aware they have a mouth, Meltzoff and Moore, 1977), and in individuals with lesions in frontal brain areas associated with mirror neuron activity (e.g. Lhermitte et al. 1986). From time to time normal individuals will unconsciously copy some basic movements, such as moving their lips as they watch someone speak. There are plenty of anecdotal and experimental examples of a ready-made copying system out there. Ramachandran pointed to skin nerves vetoing MN signals "empathise all you want, but you aren't being touched" and this veto effect disappears once the skin nerves are taken out of action. Baldissera, Cavallari, Craighero (lol) and Fadiga (2001) had a more specific suggestion. They found that the spinal cord would give the muscles which corresponded with an observed movement opposite directions...as if to cancel out what the body would have done. So if an actor moves his hand up, the observer's mirror neurons signal for his/her hand to also move up, but then the spine (closer to the muscles) tells his/her arm to move down, effectively neutralising any command at all. Bizarre!

6 participants were given a sequence of really short videos representing a number of motor acitons (right hand closing onto a ball / enlarging a rubber band / at rest). The videos showed both a "closing", "opening" and control scenario, whilst the participant's relevent nerves were monitored using electrodes. The electrodes stimulated something called the "H-reflex" which is pretty complicated beyond my understanding, but seems to be a way of testing activity in the spinal cord. By observing the change in H-reflex size over the conditions, the researchers could examine what the spinal cord was doing to some extent during those conditions. Their results showed that the spinal cord consistently and reliably activated in response to observed hand movements. In addition, the spinal cord appeared to behave in the opposite way of the oberved on-screen action (as mentioned above). The researchers suggested that the human motor system had both a cortical and spinal component which worded different from one another. The spine coordinates during an executed movement, but performs reversed actions in response to an observed movement to prevent the body replicating that movement.

Nullification through opposite response. Pretty fascinating.

It seems well established that with most behaviours there is a sense of empathy and then a system to prevent one copying or experiencing the same touch sensations. Empathy, however, is generally common to every such test. Observers feel, to some emotional extent, what the actors are experiencing. This makes me very interested not in movement (per se) but in the emotions behind the mirror neuron activity. When we observe someone doing a handstand we know we aren't doing it, but if we've ever done a handstand before we know how it feels and we can imagine it to a strong or weak degree depending on how fresh the memory is. We know what it feels like to touch ice or to experience the warmth of central heating or a hot water bottle. Every sensation brings with it some form of emotional factor, even if it is so small it can barely be described in such terms. My main overarching questions are: Does emotion have mirroring properties? If so, to what extent do we share the emotions we observe (or read about, or otherwise empathise with) and to what extent to we inhibit feeling the same way as those we observe?

Slightly more specific questions: When a human is feeling an emotion very strongly, but then is presented with an image or movie (for example) of another human expressing a powerful version of the opposite emotion, what happens? How does the mirror neuron response / experience of the emotion differ to when there is no opposite emotion observed? Are there any sex differences in perception change? Additionally, will traits such as empathy / systemising affect it? What are the neurological underpinnings of any proposed emotional mirroring system if one does exist?

The next step of my research is to work out my specific research quest to allow me to enter into this field in small, manageable steps, and how to test for it using equipment I've never even seen, let alone used.

Sex differences, social abilities and a ticking clock.

Phase two of the pre-meeting swat up. My flights are booked, my bags are packed and all I have left to do is drill this stuff into my brain. It's not a chore however. I find this stuff genuinely interesting and it will be nice to learn what directions to researchers intend to take it in, in future. It isn't often I get to look at a body of research and try to imagine what spin off studies I could create from them.


Introduction

There is a study listed in the 'selected publications' section of each researcher I have been in contact with down in Roehampton: Dr Silas, Dr Levy and Dr Holmes. I figure examining a project all three contributed to could get me a really good insight into the department. The introduction of the paper begins by mentioning mirror neuron research that I gave an overview of in my last post (good start?). Specifically it references the hypothesis and proposals made by the big names in the area (e.g. Gallese, Rizzolatti, Lacoboni) that in order for an individual to experience empathy and understanding they would need direct-matching mirror neurons. If you will recall, this is the idea that when someone observes an action that matches their mental representation of said action, they understand the purpose of that action. In the last post I mentioned research which suggested humans have seperate brain areas for representing both purpose-of-action and the physical requirements of siad action. Though there is evidence for direct-matching, Silas, Levy and Holmes did not believe there was sufficient understanding of how such a system might actually function.

The researchers planned to examine the direct-matching hypothesis in more detail, using EEG measurements. EEG (or electroencephalography... bit of a mouthful) measures the voltage of the electrical signals created in neurons. It has a heck of a lot of medication diagnostic applications, but is fairly often used in research due to the lack of invasiveness and the relative accuracy with which it shows brain activity. It even makes participants look kinda cool.

One of the more minimalistic sets of EEG equipment. Helpfully provided by the wikipedia page.

They would be studying two types of EEG measurement: Induced mu (frequencies between 8 and 13 Hz) desynchronisation and evoked readiness potentials. Now at this point I ahve absolutely no idea what either of those terms mean, so let's try and find a couple of definitions and examples... Evoked potentials are action potentials (nerve firing) that are recorded after participants are given some form of stimulus. Generally many suck trials are recorded so that the background 'noise' of other biological systems and the EEG equipment itself is cancelled out. Since the action potentials occur upon presentation of the stimulus (i.e. are time-locked), researchers can usually spot them amongst the randomly produced background distractors. This average signal rate is then analysed for "how frequently did the neurons fire during stimulus presentation?" giving an event related response (ERR). Induced mu desynchronisation measurements are based on the fact that  'mu' EEG frequencies decrease when body movement is imagined, observed or actually executed. When the ERR readings and random background noise are substracted from these measurement, one has the total increase or decrease in power of a task that cannot be explained by either of those variables. Man, this is a very difficult topic for those of us who have never used EEG equipment, but I think I have the gist of it.

Silas et al. mentions that in austistic individuals, little or no mu desynchronisation has been shown in autistic participants when compared to control groups (e.g. Ramachandran and Pineda, 2008). What this suggested was that autistic people may have malfunctioning direct-matching system, preventing them from understanding behaviours or developing empathy; two major features of autistic individuals. This viewpoint is termed the 'Broken mirror theory' in reference to mirror neurons. The causes and characteristics of autism are still a very controversial issue and this is certainly not an unargued theory, but it is what Silas et al. went on in this study. Autistic individuals have been found to have normal, higher, lower or no change in mu suppression than controls in a wide range of studies, leaving me with questions as to whether mu desynchronisation can reliably be used to measure mirror neuron activity. If it can, there are certainly other factors involved, but the broken mirror theory does not strive to accound for 100% of autistic symptoms.

Cheng et al. (2008) examined 20 males and 20 females for mirror neuron activity via mu-de measurements whilst they observed either hand movements or a moving dot. The hand movements were a mix of male-typicaly, female-typical and androgynous actions. Females, it was found, displayed significantly stronger mu suppression than male during the hand movements phase, but there was no real difference when watching the dot. The participants had been asked to fill out a survey intended to test empathy, called the "interpersonal reactivity index" before the main experiment (you can download and view the index here: http://www.suu.edu/faculty/barney/PSY3430/Factor%20Analysis%20Example/INTERPERSONAL%20REACTIVITY%20INDEX.doc ). This allowed the researchers to note that individuals who scored higher in empathy had increased mu suppression, whilst those who scored high in 'systemizing' on the opposite side of the spectrum had lower mu suppression. This view fit snugly with the extreme male brain hypothesis of autism, which suggests that the disorder is essentially an expression of exteme 'maleness'. Exaggerated versions of qualities males possess more typically than females. Wikipedia's introduction does a good job of explaining systemising:

" (defined as the drive to analyse or construct a system). A system in turn is defined as anything that follows rules, key classes of systems including mechanical systems, natural systems, abstract systems, and collectible systems. Rules in turn are defined as repeating, lawful patterns."

Systemising was negatively correlated (as systemising went up) with the degree of mu suppression (the degree went down). Not all measures of empathy were correlated in the alternative (or indeed any) direction with mu suppression, which would be expected in both broken mirror and extreme male brain theories. Research is all over the place in this area due to different methods used in each study. Puzzo et al. (2008) found no sex differences, but found the same correlation between systemising and suppression. They used the autism quotient survey to determine that those with higher scores did not display any significant suppression at, weirdly, beta frequencies rather than mu frequencies. This is all too hard to understand for a beginner in the scene! Time to simplify:

Yeah, I'm just certain empathy bear has tried learning about EEGs before. Bears go around doing that all the time. Git.

In Silas et al's study, both induced and evoked measures of a known mirror neuron system were recorded whilst participants acted and observed simple tasks prompted by a tone. ERRs had been shown to more clearly display potential mirroring between action and observation more than mu suppression, and this was the first study to use them in the study of sex differences in this area. Additionally they asked participants to self report on topics of social cognition, The researchers aim was to test whether their EEG data would reflect the self reports of social cognition. Essentially this was an attempt to gain a better insight into sex and individual differences in direct-matching systems than the confused previous findings. If direct-matching and human social understanding / empathy are strongly linked, then there should be a strong covariance.

Methods

33 participants' data were used in the final analysis; 16 males and 17 females with a mean age of 25.7 years. They were seated individually in a dimly lit room in front of a computer monitor. Baseline mu states were recorded by having participants watch a 2d circle bounce around the monitor (giving the researchers readings for how the participants' mirror neurons reacted upon seeing non biological movement). For the main condition participants took part in both an executing and observation section. For execution, they were asked to fixate their eyes upon an LED sitting just above a button which they would need to press to give a response. If a high tone was given they would press the button on the right. A low tone meant the button on the left. For observation participants were again asked to fixate their eyes on the LED but were now sitting 45 degrees to the left of the experimenter who would be pressing the buttons similarly to in the execution phase. Three blocks of each type of trial were presented to each participant alternating between the two types. Each block consisted of 80 random high or low tones. In addition, participants completed the previously mentioned index, a systemising quotient and an empathy quotient. EEG data was taken for each participant in some currently unfathomable way, during testing. Some equally baffling form of electrical magic happened to make the data understandable, and there we have it. A results section! I really wish I had managed to find experience with an EEG before going to this meeting, but I doubt they'll expect that technical know-how from a novice.

Results... but mainly discussion of results since EEG data is a bit tricky at present

To begin with, check out this shiny image of a typical participant's EEG output:

One day I will learn all about EEG, come back to this post and explain it 100 times better than I currently have.

You can see how the same brain areas are involved in both the observation and executing of the button presses, though observing has a less wide-reaching response. This fits with studies of mirror neurons showing that only a certain percentage of neurons in movement-processing areas are mirrors.

Ok, the findings showed that participants' scores on the social cognitive scales were in the expected directions as defined by sex (this is, females scored higher on empathy and males scored higher on systemising). As expected females showed stronger mu suppression than males during observation, but mu suppression did not correlate consistently with scores on the social cognitive scales. This fits quite comfortably with the confused mess that is the collective findings of all previous research in the area. During execution there was no real sex difference in suppression. The data thus supports Cheng et al's findings that there is a reliable sex difference in mirror neuron response, with females showing a higher response rate. It does not, however, support the notion that social abilities are linked to mirror neuron abilities. It must be remembered however that the method used to determine varying levels of those abilities was self reporting; a method fraught with validity problems. Evoked response potentials did not correlate with self-reported measures of social cognitive abilities either. Nor did the ERPs show that females had more mirror neuron activity than males; the opposite was in fact true. The lack of consistency in result did not support a unified underlying system of mirror neuron response.

It has been suggested by previous researchs that induced measurements may reflect transmission of sensori-motor data, whilst evoked measurements might reflect 'readiness to respond'; two fundamentally different functions. There were sex differences in opposite directions for each type of mirroring (if there are indeed two types, this is now going into advanced level), and no reliable correlation between either of them and social abilities.

My tentative conclusion of the meat of this study, is that the very complicated subject of mirror neurons is entering into a stage where they are understood enough to provoke researchers to try to learn more, but not well understood enough for their implications of human behaviour to be pinpointed. I will find out what stage Silas, Holmes and Levy are at in developing further research into the phenomenon, and who knows perhaps I will be of help in the near future. I'm encouraged by the fact my interest is blossoming and that there are so many important questions yet to be answered about a thrilling subject. Many topics in psychology are so over-researched (the value of eye-witness testimony, for one) that I would struggle to think of a single study I could do to ehance world knowledge.

This has been a struggle to read, I can only imagine. If you get this far thank you for sticking with it. If you have any questions please ask me so I can improve my understanding of the area by filling in some gaps in both our knowledge. Tomorrow I will focus on learning a little more about anxiety, emotion, language and the brain.

Mirror neurons|snoruen rorriM

For my meeting in Roehampton on Thursday, It would be a large advantage for me to learn a little more about mirror neurons and the big research in that area. Once I've informed myself, I should also check out some articles written by the researchers I will be seeing, and the research aims of their department in general. The purpose of today's blog post is to help me explain my findings to myself, via explaining them to you. A popular adage amongst teachers is "If you can teach other people the material, then you understand it well enough." In favour of that, if you read through this and have any questions, please get back to with them before thursday so I can clarify or study the answers I should give.

This was taken in my old flat, but it gives you an impression of my current work station. This is how research should be done!

Discovery and Basics of Mirror Neurons:

A quick wikipedia examination tells us that mirror neurons were discovered in an experiment in 1992 that was designed to examine what occured in an area of the brain called the premotor cortex (part of the frontal lobe, just to the front of the motor cortex. Apparantly it is very involved in the ability to guide one's movement using one's senses). This study is available as an open and free .pdf so I've had a brief read through.

Pellegrino et al. attached micro-electrodes to individual neurons in the premotor cortex of macaque monkeys, which both recorded neural activity in those neurons and 'microstimulated them'. The researchers carried out a previous study in which they outlined the methods in more detail so I am unaware of the exact procedure for now. Objects of different sizes and shapes were displayed in different positions in the visual field of the macaque. This was repeated until the researchers were certain that the neurons they were recording became active during the macaque's hand movements. Presumably it must be fairly tricky to accurately find a 'hand movement' neuron. I certainly wouldn't know where to look. For the main experiment, Pellegrino et al. presented the monkey with a large box, the front 'door' of which was made up of a one way mirror. There were different shapes inside the box which could not be seen by the monkey. The monkey was directed to press two little plates with thumb and index finger. If the monkey held on for 1.5 secondsish, then the door would open and the monkey could reach the shapes...which very pleasantly had food underneath them.The researchers used a movement recording system called ELITE to compare actual hand movements with the readings coming from the microelectrodes.

As somewhat of a fortunate surprise, the researchers noticed that when they themselves performed actions in front of the monkey, the monkey's recorded neurons would 'fire' (have an action potential, which is essentially a term for a burst of neural activity). The researchers might be presenting food the the monkey, picking it up, moving it, playing with it. Action potentials occured whenever they did any sort of hand movements within the macaque's visual field.

Macaca nemestrina, used in the experiment

The results were pretty astonishing. The microelectrodes picked up very similar readings for when the macaque made certain hand movements, and when it watched the researchers make them. This did not occur for all types of movement. For example moving hands alone would not trigger the neurons, but grasping food with them would. Grasping the same food with tweezers would not. What this suggested was that goal orientated movements which the monkey could physically copy (i.e. use hands to pick up food) were represented in some way inside the premotor cortex. The monkey seemed to be imagining making the same movement. Watch your friend opening a can of coke or punching something, using a needle and thread or picking up a really nice looking burger... at this point it seems like your brain may actually 'code' for the movement even though it isn't you making it. This has many implications in how we are able to learn new physical skills from one another.

In 1996 many of the same researchers from the same group as the previous experiment (primary researcher: Vittorio Gallese) performed a more detailed examination of the mirror neuron phenomenon. They recorded 532 neurons in "area F5" of the brain. I'm not sure of the details of this brain area, but it is referred to as a position where mirror neurons have regularly been found to exist. Research has shown that F5 is related to goal-directed hand movements but there is also some activity when individuals view 3D objects. Of the 532 neurons in each of the two macaques they used, 92 of them were mirror neurons. These neurons were discernable because they fired both when the macaques performed hand actions, and when similar actions were performed by the experimenter. Again, the researchers found that for a mirror neuron to fire, there needed to be an interaction between actor and object. Watching the actor perform an action, or just watching the object, did not produce any firing.  In addition, the mirror neurons were often incredibly precise. Around 30% of them required the macaque view both the basic movement (gripping) and a specific way in which that action was performed (gripping using certain fingers on certain positions). Gallese et al. suggested that the mirror neuron system formed a system which allowed the monkeys to learn physical movements from observation. In other words, allowed them to copy. The argument has been presented that the mirror neurons were really just preparing the monkeys to make the same movement. This was rejected by the finding that the mirror neuron response decreased when the experimenter moved the object within the reach of the monkeys, and only increased again when the monkeys physically grasped the object.

Mirror Neurons and Social Cognition

In a later experiment, Ferrari et al. found strong evidence showing mirror neurons also activated in response to observing mouth movements. Mostly those that were related to eating behaviours. Very relevent to social cognition was the finding that the most reliable way to fire some mirror neurons in the monkey subjects was to have them observe communicative gestures and lip movements. This experiment opened my eyes to the importance of this F5 area in the control of both mouth and hand movements.Rizzolatti et al. pointed out that understanding acitons made by others is essential to social interaction. If mirror neurons exist to aid action understanding, that it may work like this: The observer has a mental representation of what happens when he, say, moves his arm to grab food. He understand that his hand will move from point A to point B. When an observed action matches this mental representation then the observer understands the action.

Showing how readily a newborn macaque will copy the mouth actions of an experimenter. Human babies are also very capable of this from birth, suggesting that there is a ready-built system for copying rather than a simple learned ability. Perhaps mirror neurons are the answer.

35% of mouth motor neurons fired when the macaques in the experiment observed mouth actions. These were thus considered mirror neurons specific to the mouth area. 85% of these fired in response to eating behaviour. 15% to communicative behaviour. The experiment was controlled so that movements made by the macaques during observations were accounted for and so not mistaken for mirror neuron firing. Again, interaction between actor's lips and an object was necessary for mirror neurons to fire, bringing further support of them action as facilitators of action understanding. In humans an fMRI study by Buccino et al. showed evidence for mirror neurons activated in response to watching chewing and biting movements. This activation occurs in area 44 of the human brain, which has been considered the homologue of monkey's area F5 (e.g. petrides and Pandya, 1994).

Communicative mirron neurons were different. They didn't necessitate goal orientated lip movements to fire. Lip-smacking, tongue protrusions etc are not directed towards any object. It is possible that the actions prompted the macaques to imagine the result of that sort of expression when made in the presence of another macaque. These mirron neurons barely fired in response to any ingestive action, suggesting that they are specific to communication to some extent. There is a methodological problem in examining communicative behaviour in an ecologically valid setting, so many questions remain in this area.

An example of the sort of data collected from the macaque experiments. You can see that the strongest mirror neuron response was to interactions between the experimenter and the food. The firing was even stronger when the experimenter used a tool, this will be explained under the last section of this blog entry.

Mirror Neurons in Humans?

fMRI machines are the best friend of researchers looking to examine neural activity in humans. Attaching electrodes to the neurons is a horribly invasive procedure and I still very much dislike the thought of that method being used on macaques, but you know. Science -.-. fMRI scans scan the changes in blood flow throughout the central nervous systems (brain and spinal cord) of anyone in the machine. More blood flow essentially means more activity in the brain. By giving participants tasks or objects to observe whilst they are in the machine, researchers can gain an understanding of what brain areas are involved in what processes.

You've all seen pictures like this before. The coloured areas represent different levels of higher-than-normal blood flow. It clearly displays active areas in the brain.

Lacoboni et al. write that imitation is crucial for human development and practise of communication and social skills. In order to examine the neurological underpinnings of our ability to imitate, the researchers set up an experiment where participants were asked to observe and then copy simple finger movements...whilst in an fMRI machine. Human neonates can copy facial expressions and really basic gestures within a few hours of birth. If you don't believe me, check out this baby of a video!

Direct link: http://www.youtube.com/watch?v=k2YdkQ1G5QI

Remember I spoke about matching an observed action to a mental representation? That is called the "direct matching hypothesis". Lacoboni et al were testing for it. If mirror neurons had a 'matching mechanism' then they should 'fire' more prominently when the participants observed someone actually moving their finger, rather than some symbollic cues. See how that works? If mirror neurons work by activating whenever reality matches imagination, then observing a note saying "Move finger two squares to the right" should not activate mirror neurons.

Indeed, this was the case. Signal intensity was significantly stronger for the direct match observations than for symbols, though there was still some firing in the same brain regions (basically: area 44 or Broca's area. Check it out on wikipedia and you will notice it is described as being primarily linked to speech production. Hence the afflication Broca's Aphasia). Lacoboni et al. saw there being good reason for mirror neurons to be present in that area, seeing as it is the human version of F5 and is known to represent 'distal' movement. Proximal: Area where appendages join the body. Distal: Area furthest from where the appendages join the body. Aren't we learning a lot of terminology today? Additionally, Broca's area is all about speech production, and Lacoboni et al suggests that as imitation is crucial in language acquisition, it makes sense that there would be some neural propensity towards imitation learning there. I'm not too sure about that one after having puzzled out some of Vera Kempe's perspectives on pre-lingual language development, but we'll see. Fourth and last, speech production not only involves learning to words, but learning how to pronounce them physically possibly via direct matching.

Other areas are involved, according to the researchers, but the reasoning behind them is less well understood. Previous research had used PET scans to observe activity in human brains whilst they watched pantomimes involving complex movements. Participants were asked either to memorise and attempt to copy the movements, or try to work out their purpose. In the first situation the parietal lobe was activated, mainly on the right. The latter sitation saw more activation in the left inferior frontal lobe (same as above). Their interpretation of this research was that a mental representation of the final position and the steps necessary to get to it may have been building in the parietal lobe whilst watching. Broca's area, in contrast, builds a representation of the goal of the body movements but not the actual physical movements needed to arrive there. This hypothesis totally works with F5 areas in Macaques not activating unless there is interaction between actor and object (goal orientated behaviour). A whole wealth of research shows the lesions to these areas causes problems with action understanding, direct matching and other abilities described above. Mirror neurons, although only indirectly observable in humans, seem very clearly to exist in a similar way to macaques.

Scratch that above sentence, humans have more recently been implanted with electordes for direct examination of the existance of mirror neurons. This was in 2010 by Mukamel et al. (I love how many of these articles are free and available to view just with a quick googling! In university so many of the brilliant articles I needed had to be paid for and werent under Abertay's subscription. Also every researcher in this whole area seems to be Italian so far...). They examined 1177 neurons in relevent areas whilst the participants hooked up to the electrodes performed or watched hand graspig and facial expressions. These folks were patients already being treated with invasive surgery so ethically it was cool. That's a relief. Many of these neurons acted as mirror neurons would be expected to act, but some others actually activated during activation but were inhibited during observation so that activation was significantly lower than normal in those neurons. An interesting finding. The rest of the results were consistent with the indirect human research using fMRIs and with the animal research.

How and when do mirror neurons develop in human infants?

So we seem pretty sure that mirror neurons exist, and that they aid in learning how to speak, function in social situations and perform certain types of actions. As we spoke about before, newborn humans have shown the ability to imitate just hours after birth. In an article entitled "Where do mirror neurons come from?" Cecilia Heyes explores possible models of their development. There are no current studies showing mirror neurons existing in newborn infants (you can see why) but many researchers argue that being able to imitate = mirror neurons. A brilliant review of facial expression mimicry in neonates actually finds that the vast majority of reports of this involve just the tongue-poking out action (Anisfeld, 1996). More recently, a very similar finding was discovered for lip-smacking and tongue protrusions in rhesus macaques. If these results are really the trend, then the mimicry of neonates could be put down to more basic copying based on the tongue, not something so complex as mirror neurons.

Some examples of how similar the human and monkey behaviours could be in some of the experiments described in this post. The guy on the right reminds me of a different actor in each photo.

Studies, like one by Ferrari et al. (2005), have shown that mirror neuron responses depend to some extent on life experience. Ferrari et el. gave monkeys opportunities to see human experimenters using tools (a stick / pliers) to pick up or manipulate food. When examined for mirror neuron activity whilst observing these experimenters after several trials, the monkeys exhibited significant firing (as in the figure showing graph output further up the page). This is contrary to the earliest research with macaques that showed no mirror neuron firing when the monkeys observed the experimenter picking up food with tweezers. Experience with tool use had altered the firing responses of the mirror neurons. Haslinger et al. (2005) discovered more mirror neuron activity in human pianists when observing someone playing the piano, then in human non-pianists. Similar results were found in an experiment involving classical ballet dancers (Calvo-Merino et al. 2005). How could mirror neurons be present and functioning at birth if experience in both acting and observing has such a profound effect? Heyes added that certian combinations of sensory input and motor action could actually produce counter-mirroring responses. Catmur et al. (2008) gave various examples of this, including how after specific confusing training, observing movements in the index finger could actually provoke mirror neuron activity in brain areas controlling the pinky finger. The observation of stimuli A had provoked the mental representation of action B in participants who had the experience of the training exercises designed to misdirect.

Heyes concluded that mirror neurons reflect the behaviours and observations we have experienced. This is essential associative learning, which is a well documented phenomenon that has been observed in human and most non human species. As a result, might it be expected that most animal species would have mirror neurons? Heyes goes so far as to suggest that it might be possible to create mirror neurons in animals that wouldn't otherwise have them, by giving them training to associate an interacting behaviour with an itnernal representation. Bizarre thought. Research, it is said in her article, on the effects of mirror neurons is just beginning. If they are produced as a result of associative learning then it does not make sense that they are responsible for prompting humanity to jump forward in its incredible social learning and develop as a species. She believes it is more likely that human sociality boths develops and is developed by mirror neurons. A product and a producer of social interaction. Hmmm. Leaves a lot of questions unanswered. I'm not sure what my perspective is on this yet.

That is enough of an introduction to the topic for now I reckon! I will do more study tomorrow on more specific articles that will or will not be related to mirror neuron research. I also plan to review my work on anxiety (which I mainly directed towards panic disorder) and re-examine the assessments for which I studied Libet and Ramachandran's work. With enough time I will take a quick glance over social brain hypothesis stuff again, and hope that that will be enough work to provide a good impression at this meeting. Good start tonight, I reckon :).

Hopefully I wont Roe the day I went to this meeting.

I tossed the idea of writing a new blog back and forth before settling on keeping Radjeworks. It's been a hell of a long time since I wrote anything of any depth, and a lot of it shimmers with the sort of innaccuracies you only notice after working your face off studying them for exam season and dissertation. It stands as a testament to the learning process though, and there are a few articles scattered within that talk a little about my own mini-adventures. Consider everything after the last post (in Feb 2011) to be of the past, and we'll move on to some exciting adventures in the upcoming few days.

It seems odd, considering how all-consuming 4th year was at the time, that I can sum up the past few months in a few paragraphs. The rest of February, March and April were dominated by the 100% examination modules and my honours project. My friends and classmates in Dundee will remember (with varying degrees of distaste) what it felt like to have every conversation eventually lead back to coursework and the impending doom of assessment. Procrastination became a way of life for me, not because I was a lazy student but because I could no longer feel entirely guilt-free when not doing coursework. Perhaps because of that, I pushed all of my focus into my honours project. If you're interested; the final title was "The application of models of sexual selection to between sex competition". It gave me an excuse to go into way more detail about evolutionary perspectives of human behaviours rather than get stuck looking at the less thrilling 'social psychological' aspects.

Along with Advanced cognitive psychology, advanced forensic psychology and education and health, I passed my last year with flying colours and accepted a 1st class degree with pride and a great deal of relief. Graduation was an absolutely wonderful experience. Strawberry tarts, champagne, garden parties on warm summer afternoons, the incredible ceremonies and most importantly, the friends and their oh so very proud families. Standing beside my good friends Gavin (1st class in computing) and Paul (1st class in Mechanical Engineering) was a special moment. All from the same year, same school, same town.

So. What's next?

Summer was spent in a really well decorated and plush limbo in which I had a really great time. I moved out of my flat in Dundee... I miss it, but it was definitely time to leave... to come back to Dunfermline to live with my parents. The PhD / Research assistant job hunt became all the more fervent for the loss of independence, but my parents have always been understanding, fair and nice to be around. We havent had to work too hard to keep the peace :). During the first part of summer I stayed with Paul and Katie so I could carry on at the library and save like crazy for the gap between employment and employment. Paul and I got through a lot of ace multiplayer games and I haven't had a good cup of tea since the last one Katie made me. Sigh. I had forgotten the good parts of living with flatmates. Eventually though they moved back to their respective homes also, and I had to leave the library behind. The ladies at Douglas made sure I departed in style with a case of beer and loads of pastries!

Little bit of free publicity for Dean Koontz here.

After coming home, there wasn't much to do besides apply for work/ PhDs and spend time with equally restless friends. Edinburgh zoo, Dunfermline Glen to feed the squirrels, Glasgow, Stirling to do the tourist thing. Lots of visits to pubs and playing games, all the time taking breathers to apply for a position here or there. The process has been one of rejection and a lack of feedback thus far, but it was never going to be simple so my moral hasn't dropped too much. At the beginning of August I used a chunk of my savings to go on a great venture to France with parents, then back to Norway to see Mia'n'co. We spent most of the time in Mia's Cabin just across the border in Sweden, where I got to meet such a fantastic group of their friends. We went canoing and swimming in the nearby lake, had campfires and played an overwhelming amount of yahtzee. It was here I made friends with Torun, who will feature later on in this summary!

Every one of these people is an individual and compelling argument for a huge bridge connecting Scotland and Norway.

Now that Autumn has begun, things have changed. The focus is now very strongly on the application process and a sort of terror-based curiosity about where I will end up and what I will be doing. After a meeting with my Ex-tutor Fhionna, I started volunteering with Vera, the tutor who taught the communication part of communication and consciousness; and basically all of the language stuff we've done in the past 4 years. More recently Vera approached me to act as a casual paid research assistant on a different project. One of the people I met during my duties in the volunteer research, Kenny McAlpine, expressed interest in having me give a small seminar to some of his 4th year students about the research process. In short, I've begun adding bits and pieces to my CV through a combination of genuine interest and luck. My friends have, in the main, already left limbo and are well on their way to future success. I have to congratulate them on how readily their hard work in Uni has paid off! 

Only a couple of days ago, I received a reply to an email I sent to a tutor in The University of Roehampton enquiring about some advertised PhD positions. They were the type of advertisements that gave you free reign to design your own project proposal, but hinted very strongly that the best path to success in the process would be to get in touch with some of the potential supervisors and see what they suggest. My attention was grabbed by the department of psychology's access to an fMRI machine, and the centre for research in cognition, emotion and interaction (CRICEI)'s focus on combining neurological data with psychological phenomenon. To really top it off, these findings are often applied to areas such as anxiety and social behaviours. The tutor expressed an interest in my explanations of my motivation for applying for the PhDs there, and invited me to come meet with him and a co-worker who was also keen to meet me. I was thrilled at a chance to discuss their work and to research it well enough to discuss potential projects leading from it. As a result, I have booked flights and will be heading down to Roehampton this coming week! Torun, you will recall me mentioning a couple paragraphs above, has kindly offered me a place to stay for the couple of days I will be down south. I can't wait to see her, and to be getting on a plane again. 

This weekend is dedicated to researching the work of Dr Jon Silas, Dr Joe Levy and many of the big names in the mirror neuron area as suggested to me by my old tutor, Clare Cunningham. If this meeting goes well, it could be a real advantage to my eventual application, and Roehampton is definitely a really good place to end up. I believe my blog would be a good way to write up my train of thought as I gain more insight into the complicated subjects, so keep your eyes peeled for more in the upcoming days. Maybe even hours.