Sorry for the brief(?) hiatus. I have left my day job to start a venture and so am a bit preoccupied. Hopefully, the mouse trap should benefit from the new arrangements.
Today I would like to highlight a recent study from MIT that once again highlighted the fact that the same brain mechanisms are used for envisaging the future as are used for reminiscing about the past. The study was performed on rats and found that the rats sort of replayed their day-time navigational memories while they were dreaming. This in itself is not a new news and has been known for a long time; what they found additionally is that the rats also , sort of replayed the navigational memories/ alternatives in their head at a faster rate, to sort of think and plan ahead. This use of replaying the traces to think ahead to me is very important and cements the role of default netwrok in remebering the poast and envisaging the future.
When a rat moves through a maze, certain neurons called “place cells,” which respond to the animal’s physical environment, fire in patterns and sequences unique to different locations. By looking at the patterns of firing cells, researchers can tell which part of the maze the animal is running.
While the rat is awake but standing still in the maze, its neurons fire in the same pattern of activity that occurred while it was running. The mental replay of sequences of the animals’ experience occurs in both forward and reverse time order.
“This may be the rat equivalent of ‘thinking,'” Wilson said. “This thinking process looks very much like the reactivation of memory that we see during non-REM dream states, consisting of bursts of time-compressed memory sequences lasting a fraction of a second.
“So, thinking and dreaming may share the same memory reactivation mechanisms,” he said.
“This study brings together concepts related to thought, memory and dreams that all potentially arise from a unified mechanism rooted in the hippocampus,” said co-author Fabian Kloosterman, senior postdoctoral associate.
The team’s results show that long experiences, which in reality could have taken tens of seconds or minutes, are replayed in only a fraction of a second. To do this, the brain links together smaller pieces to construct the memory of the long experience.
The researchers speculated that this strategy could help different areas of the brain share information – and deal with multiple memories that may share content – in a flexible and efficient way. “These results suggest that extended replay is composed of chains of shorter subsequences, which may reflect a strategy for the storage and flexible expression of memories of prolonged experience,” Wilson said.
To me this seals the fate of hippocampus as not just necessary for formation of new memories, but also for novel future-oriented thoughts and imaginations.
This article continues my series on major conscious and unconscious processes in the brain. In my last two posts I have talked about 8 major unconscious processes in the brain viz sensory, motor, learning , affective, cognitive (deliberative), modelling, communications and attentive systems. Today, I will not talk about brain in particular, but will approach the problem from a slightly different problem domain- that of modelling/implementing an artificial brain/ mind.
I am a computer scientist, so am vaguely aware of the varied approaches used to model/implement the brain. Many of these use computers , though not every approach assumes that the brain is a computer.
Before continuing I would briefly like to digress and link to one of my earlier posts regarding the different traditions of psychological research in personality and how I think they fit an evolutionary stage model . That may serve as a background to the type of sweeping analysis and genralisation that I am going to do. To be fair it is also important to recall an Indian parable of how when asked to describe an elephant by a few blind man each described what he could lay his hands on and thus provided a partial and incorrect picture of the elephant. Some one who grabbed the tail, described it as snake-like and so forth.
With that in mind let us look at the major approaches to modelling/mplementing the brain/intelligence/mind. Also remember that I am most interested in unconscious brain processes till now and sincerely believe that all the unconscious processes can, and will be successfully implemented in machines. I do not believe machines will become sentient (at least any time soon), but that question is for another day.
So, with due thanks to @wildcat2030, I came across this book today and could immediately see how the different major approaches to artificial robot brains are heavily influenced (and follow) the evolutionary first five stages and the first five unconscious processes in the brain.
The book in question is ‘Robot Brains: Circuits and Systems for Conscious Machines’ by Pentti O. Haikonen and although he is most interested in conscious machines I will restrict myself to intelligent but unconscious machines/robots.
The first chapter of the book (which has made to my reading list) is available at Wiley site in its entirety and I quote extensively from there:
Presently there are five main approaches to the modelling of cognition that could be used for the development of cognitive machines: the computational approach (artificial intelligence, AI), the artificial neural networks approach, the dynamical systems approach, the quantum approach and the cognitive approach. Neurobiological approaches exist, but these may be better suited for the eventual explanation of the workings of the biological brain.
The computational approach (also known as artificial intelligence, AI) towards thinking machines was initially worded by Turing (1950). A machine would be thinking if the results of the computation were indistinguishable from the results of human thinking. Later on Newell and Simon (1976) presented their Physical Symbol System Hypothesis, which maintained that general intelligent action can be achieved by a physical symbol system and that this system has all the necessary and sufficient means for this purpose. A physical symbol system was here the computer that operates with symbols (binary words) and attached rules that stipulate which symbols are to follow others. Newell and Simon believed that the computer would be able to reproduce human-like general intelligence, a feat that still remains to be seen. However, they realized that this hypothesis was only an empirical generalization and not a theorem that could be formally proven. Very little in the way of empirical proof for this hypothesis exists even today and in the 1970s the situation was not better. Therefore Newell and Simon pretended to see other kinds of proof that were in those days readily available. They proposed that the principal body of evidence for the symbol system hypothesis was negative evidence, namely the absence of specific competing hypotheses; how else could intelligent activity be accomplished by man or machine? However, the absence of evidence is by no means any evidence of absence. This kind of ‘proof by ignorance’ is too often available in large quantities, yet it is not a logically valid argument. Nevertheless, this issue has not yet been formally settled in one way or another. Today’s positive evidence is that it is possible to create world-class chess-playing programs and these can be called ‘artificial intelligence’. The negative evidence is that it appears to be next to impossible to create real general intelligence via preprogrammed commands and computations.
The original computational approach can be criticized for the lack of a cognitive foundation. Some recent approaches have tried to remedy this and consider systems that integrate the processes of perception, reaction, deliberation and reasoning (Franklin, 1995, 2003; Sloman, 2000). There is another argument against the computational view of the brain. It is known that the human brain is slow, yet it is possible to learn to play tennis and other activities that require instant responses. Computations take time. Tennis playing and the like would call for the fastest computers in existence. How could the slow brain manage this if it were to execute computations?
The artificial neural networks approach, also known as connectionism, had its beginnings in the early 1940s when McCulloch and Pitts (1943) proposed that the brain cells, neurons, could be modelled by a simple electronic circuit. This circuit would receive a number of signals, multiply their intensities by the so-called synaptic weight values and sum these modified values together. The circuit would give an output signal if the sum value exceeded a given threshold. It was realized that these artificial neurons could learn and execute basic logic operations if their synaptic weight values were adjusted properly. If these artificial neurons were realized as hardware circuits then no programs would be necessary and biologically plausible artificial replicas of the brain might be possible. Also, neural networks operate in parallel, doing many things simultaneously. Thus the overall operational speed could be fast even if the individual neurons were slow. However, problems with artificial neural learning led to complicated statistical learning algorithms, ones that could best be implemented as computer programs. Many of today’s artificial neural networks are statistical pattern recognition and classification circuits. Therefore they are rather removed from their original biologically inspired idea. Cognition is not mere classification and the human brain is hardly a computer that executes complicated synaptic weight-adjusting algorithms.
The human brain has some 10 to the power of 11 neurons and each neuron may have tens of thousands of synaptic inputs and input weights. Many artificial neural networks learn by tweaking the synaptic weight values against each other when thousands of training examples are presented. Where in the brain would reside the computing process that would execute synaptic weight adjusting algorithms? Where would these algorithms have come from? The evolutionary feasibility of these kinds of algorithms can be seriously doubted. Complicated algorithms do not evolve via trial and error either. Moreover, humans are able to learn with a few examples only, instead of having training sessions with thousands or hundreds of thousands of examples. It is obvious that the mainstream neural networks approach is not a very plausible candidate for machine cognition although the human brain is a neural network.
Dynamical systems were proposed as a model for cognition by Ashby (1952) already in the 1950s and have been developed further by contemporary researchers (for example Thelen and Smith, 1994; Gelder, 1998, 1999; Port, 2000; Wallace, 2005). According to this approach the brain is considered as a complex system with dynamical interactions with its environment. Gelder and Port (1995) define a dynamical system as a set of quantitative variables, which change simultaneously and interdependently over quantitative time in accordance with some set of equations. Obviously the brain is indeed a large system of neuron activity variables that change over time. Accordingly the brain can be modelled as a dynamical system if the neuron activity can be quantified and if a suitable set of, say, differential equations can be formulated. The dynamical hypothesis sees the brain as comparable to analog feedback control systems with continuous parameter values. No inner representations are assumed or even accepted. However, the dynamical systems approach seems to have problems in explaining phenomena like ‘inner speech’. A would-be designer of an artificial brain would find it difficult to see what kind of system dynamics would be necessary for a specific linguistically expressed thought. The dynamical systems approach has been criticized, for instance by Eliasmith (1996, 1997), who argues that the low dimensional systems of differential equations, which must rely on collective parameters, do not model cognition easily and the dynamicists have a difficult time keeping arbitrariness from permeating their models. Eliasmith laments that there seems to be no clear ways of justifying parameter settings, choosing equations, interpreting data or creating system boundaries. Furthermore, the collective parameter models make the interpretation of the dynamic system’s behaviour difficult, as it is not easy to see or determine the meaning of any particular parameter in the model. Obviously these issues would translate into engineering problems for a designer of dynamical systems.
The quantum approach maintains that the brain is ultimately governed by quantum processes, which execute nonalgorithmic computations or act as a mediator between the brain and an assumed more-or-less immaterial ‘self’ or even ‘conscious energy field’ (for example Herbert, 1993; Hameroff, 1994; Penrose, 1989; Eccles, 1994). The quantum approach is supposed to solve problems like the apparently nonalgorithmic nature of thought, free will, the coherence of conscious experience, telepathy, telekinesis, the immortality of the soul and others. From an engineering point of view even the most practical propositions of the quantum approach are presently highly impractical in terms of actual implementation. Then there are some proposals that are hardly distinguishable from wishful fabrications of fairy tales. Here the quantum approach is not pursued.
The cognitive approach maintains that conscious machines can be built because one example already exists, namely the human brain. Therefore a cognitive machine should emulate the cognitive processes of the brain and mind, instead of merely trying to reproduce the results of the thinking processes. Accordingly the results of neurosciences and cognitive psychology should be evaluated and implemented in the design if deemed essential. However, this approach does not necessarily involve the simulation or emulation of the biological neuron as such, instead, what is to be produced is the abstracted information processing function of the neuron.
A cognitive machine would be an embodied physical entity that would interact with the environment. Cognitive robots would be obvious applications of machine cognition and there have been some early attempts towards that direction. Holland seeks to provide robots with some kind of consciousness via internal models (Holland and Goodman, 2003; Holland, 2004). Kawamura has been developing a cognitive robot with a sense of self (Kawamura, 2005; Kawamura et al., 2005). There are also others. Grand presents an experimentalist’s approach towards cognitive robots in his book (Grand, 2003).
A cognitive machine would be a complete system with processes like perception, attention, inner speech, imagination, emotions as well as pain and pleasure. Various technical approaches can be envisioned, namely indirect ones with programs, hybrid systems that combine programs and neural networks, and direct ones that are based on dedicated neural cognitive architectures. The operation of these dedicated neural cognitive architectures would combine neural, symbolic and dynamic elements.
However, the neural elements here would not be those of the traditional neural networks; no statistical learning with thousands of examples would be implied, no backpropagation or other weight-adjusting algorithms are used. Instead the networks would be associative in a way that allows the symbolic use of the neural signal arrays (vectors). The ‘symbolic’ here does not refer to the meaning-free symbol manipulation system of AI; instead it refers to the human way of using symbols with meanings. It is assumed that these cognitive machines would eventually be conscious, or at least they would reproduce most of the folk psychology hallmarks of consciousness (Haikonen, 2003a, 2005a). The engineering aspects of the direct cognitive approach are pursued in this book.
Now to me these computational approaches are all unidimensional-
- The computational approach is suited for symbol-manipulation and information-represntation and might give good results when used in systems that have mostly ‘sensory’ features like forming a mental represntation of external world, a chess game etc. Here something (stimuli from world) is represented as something else (an internal symbolic represntation).
- The Dynamical Systems approach is guided by interactions with the environment and the principles of feedback control systems and also is prone to ‘arbitrariness’ or ‘randomness’. It is perfectly suited to implement the ‘motor system‘ of brain as one of the common features is apparent unpredictability (volition) despite being deterministic (chaos theory) .
- The Neural networks or connectionsim is well suited for implementing the ‘learning system’ of the brain and we can very well see that the best neural network based systems are those that can categorize and classify things just like ‘the learning system’ of the brain does.
- The quantum approach to brain, I haven’t studied enough to comment on, but the action-tendencies of ‘affective system’ seem all too similar to the superimposed,simultaneous states that exits in a wave function before it is collapsed. Being in an affective state just means having a set of many possible related and relevant actions simultaneously activated and then perhaps one of that decided upon somehow and actualized. I’m sure that if we could ever model emotion in machine sit would have to use quantum principles of wave functions, entanglemnets etc.
- The cognitive approach, again I haven’t go a hang of yet, but it seems that the proposal is to build some design into the machine that is based on actual brain and mind implemntations. Embodiment seems important and so does emulating the information processing functions of neurons. I would stick my neck out and predict that whatever this cognitive approach is it should be best able to model the reasoning and evaluative and decision-making functions of the brain. I am reminded of the computational modelling methods, used to functionally decompose a cognitive process, and are used in cognitive science (whether symbolic or subsymbolic modelling) which again aid in decision making / reasoning (see wikipedia entry)
Overall, I would say there is room for further improvement in the way we build more intelligent machines. They could be made such that they have two models of world – one deterministic , another chaotic and use the two models simulatenously (sixth stage of modelling); then they could communicate with other machines and thus learn language (some simulation methods for language abilities do involve agents communicating with each other using arbitrary tokens and later a language developing) (seventh stage) and then they could be implemented such that they have a spotlight of attention (eighth stage) whereby some coherent systems are amplified and others suppressed. Of course all this is easier said than done, we will need at least three more major approaches to modelling and implementing brain/intelligence before we can model every major unconscious process in the brain. To model consciousness and program sentience is an uphill task from there and would definitely require a leap in our understandings/ capabilities.
Do tell me if you find the above reasonable and do believe that these major approaches to artificial brain implementation are guided and constrained by the major unconscious processes in the brain and that we can learn much about brain from the study of these artificial approaches and vice versa.
Today I plan to touch upon the topic of consciousness (from which many bloggers shy) and more broadly try to delineate what I believe are the important different conscious and unconscious processes in the brain. I will be heavily using my evolutionary stages model for this.
To clarify myself at the very start , I do not believe in a purely reactive nature of organisms; I believe that apart from reacting to stimuli/world; they also act , on their own, and are thus agents. To elaborate, I believe that neuronal groups and circuits may fire on their own and thus lead to behavior/ action. I do not claim that this firing is under voluntary/ volitional control- it may be random- the important point to note is that there is spontaneous motion.
- Sensory system: So to start with I propose that the first function/process the brain needs to develop is to sense its surroundings. This is to avoid predators/ harm in general. this sensory function of brain/sense organs may be unconscious and need not become conscious- as long as an animal can sense danger, even though it may not be aware of the danger, it can take appropriate action – a simple ‘action’ being changing its color to merge with background.
- Motor system:The second function/ process that the brain needs to develop is to have a system that enables motion/movement. This is primarily to explore its environment for food /nutrients. Preys are not going to walk in to your mouth; you have to move around and locate them. Again , this movement need not be volitional/conscious – as long as the animal moves randomly and sporadically to explore new environments, it can ‘see’ new things and eat a few. Again this ‘seeing’ may be as simple as sensing the chemical gradient in a new environmental.
- Learning system: The third function/process that the brain needs to develop is to have a system that enables learning. It is not enough to sense the environmental here-and-now. One needs to learn the contingencies in the world and remember that both in space and time. I am inclined to believe that this is primarily pavlovaion conditioning and associative learning, though I don’t rule out operant learning. Again this learning need not be conscious- one need not explicitly refer to a memory to utilize it- unconscious learning and memory of events can suffice and can drive interactions. I also believe that need for this function is primarily driven by the fact that one interacts with similar environments/con specifics/ predators/ preys and it helps to remember which environmental conditions/operant actions lead to what outcomes. This learning could be as simple as stimuli A predict stimuli B and/or that action C predicts reward D .
- Affective/ Action tendencies system .The fourth function I propose that the brain needs to develop is a system to control its motor system/ behavior by making it more in sync with its internal state. This I propose is done by a group of neurons monitoring the activity of other neurons/visceral organs and thus becoming aware (in a non-conscious sense)of the global state of the organism and of the probability that a particular neuronal group will fire in future and by thus becoming aware of the global state of the organism , by their outputs they may be able to enable one group to fire while inhibiting other groups from firing. To clarify by way of example, some neuronal groups may be responsible for movement. Another neuronal group may be receiving inputs from these as well as say input from gut that says that no movement has happened for a time and that the organism has also not eaten for a time and thus is in a ‘hungry’ state. This may prompt these neurons to fire in such a way that they send excitatory outputs to the movement related neurons and thus biasing them towards firing and thus increasing the probability that a motion will take place and perhaps the organism by indulging in exploratory behavior may be able to satisfy hunger. Of course they will inhibit other neuronal groups from firing and will themselves stop firing when appropriate motion takes place/ a prey is eaten. Again nothing of this has to be conscious- the state of the organism (like hunger) can be discerned unconsciously and the action-tendencies biasing foraging behavior also activated unconsciously- as long as the organism prefers certain behaviors over others depending on its internal state , everything works perfectly. I propose that (unconscious) affective (emotional) state and systems have emerged to fulfill exactly this need of being able to differentially activate different action-tendencies suited to the needs of the organism. I also stick my neck out and claim that the activation of a particular emotion/affective system biases our sensing also. If the organism is hungry, the food tastes (is unconsciously more vivid) better and vice versa. thus affects not only are action-tendencies , but are also, to an extent, sensing-tendencies.
- Decisional/evaluative system: the last function (for now- remember I adhere to eight stage theories- and we have just seen five brain processes in increasing hierarchy) that the brain needs to have is a system to decide / evaluate. Learning lets us predict our world as well as the consequences of our actions. Affective systems provide us some control over our behavior and over our environment- but are automatically activated by the state we are in. Something needs to make these come together such that the competition between actions triggered due to the state we are in (affective action-tendencies) and the actions that may be beneficial given the learning associated with the current stimuli/ state of the world are resolved satisfactorily. One has to balance the action and reaction ratio and the subjective versus objective interpretation/ sensation of environment. The decisional/evaluative system , I propose, does this by associating values with different external event outcomes and different internal state outcomes and by resolving the trade off between the two. This again need not be conscious- given a stimuli predicting a predator in vicinity, and the internal state of the organism as hungry, the organism may have attached more value to ‘avoid being eaten’ than to ‘finding prey’ and thus may not move, but camouflage. On the other hand , if the organisms value system is such that it prefers a hero’s death on battlefield , rather than starvation, it may move (in search of food) – again this could exist in the simplest of unicellular organisms.
Of course all of these brain processes could (and in humans indeed do) have their conscious counterparts like Perception, Volition,episodic Memory, Feelings and Deliberation/thought. That is a different story for a new blog post!
And of course one can also conceive the above in pure reductionist form as a chain below:
sense–>recognize & learn–>evaluate options and decide–>emote and activate action tendencies->execute and move.
and then one can also say that movement leads to new sensation and the above is not a chain , but a part of cycle; all that is valid, but I would sincerely request my readers to consider the possibility of spontaneous and self-driven behavior as separate from reactive motor behavior.
Till now, most of the research on learning at the molecular level or LTP/TLD has focused on classical conditioning paradigms. To my knowledge for the first time someone has started looking at whether , on the molecular level, classical conditioning , which works by associations between external stimuli, is differently encoded and implemented from operant learning , which depends on learning the reward contingencies of one’s spontaneously generated behavior.
Bjorn Brembs and colleagues have shown that the normal learning pathway implicated in classical conditioning, which involves Rugbata gene in fruit fly and works on adenylyl cyclase (AC) , is not involved in pure operant learning; rather pure operant learning is mediated by Protein Kinase C (PKC) pathways. This is not only a path breaking discovery , as it cleary shows the double dissociation showing genetically mutant flies, it is also a marvelous example fo how a beautiful experimental setup was convened to separate and remove the classical conditioning effects from normal operant learning and generate a pure operant learning procedure. You can read more about the procedure on Bjorn Brembs site and he also maintains a very good blog, so check that out too.
Here is the abstract of the article and the full article is available at the Bjorn Brembs site.
Learning about relationships between stimuli (i.e., classical conditioning ) and learning about consequences of one’s own behavior (i.e., operant conditioning ) constitute the major part of our predictive understanding of the world. Since these forms of learning were recognized as two separate types 80 years ago , a recurrent concern has been the
issue of whether one biological process can account for both of them . Today, we know the anatomical structures required for successful learning in several different paradigms, e.g., operant and classical processes can be localized to different brain regions in rodents  and an identified neuron in Aplysia shows opposite biophysical changes after operant and classical training, respectively. We also know to some detail the molecular mechanisms underlying some forms of learning and memory consolidation. However, it is not known whether operant and classical learning can be distinguished at the molecular level. Therefore, we investigated whether genetic manipulations could differentiate between operant and classical learning in dorsophila. We found a double dissociation of protein kinase C and adenylyl cyclase on operant and classical learning. Moreover, the two learning systems interacted hierarchically such that classical predictors were learned preferentially over operant predictors.
Do take a look at the paper and the experimental setup and lets hope that more focus on operant learning would be the focus from now on and would lead to a paradigmatic shift in molecular neuroscience with operant conditioning results more applicable to humans than classical conditioning results, in my opinion.
B BREMBS, W PLENDL (2008). Double Dissociation of PKC and AC Manipulations on Operant and Classical Learning in Drosophila Current Biology, 18 (15), 1168-1171 DOI: 10.1016/j.cub.2008.07.041
I had speculated in one of my earlier posts that Glutamate , GABA, Glycine and aspartate may be involved in classical conditioning / avoidance learning. To quote:
That is it for now; I hope to back up these claims, and extend this to the rest of the 3 traits too in the near future. Some things I am toying with is either classical conditioning and avoidance learning on these higher levels; or behavior remembering (as opposed to learning) at these higher levels. Also other neurotransmitter systems like gluatamete, glycine, GABA and aspartate may be active at the higher levels. Also neuro peptides too are broadly classified in five groups so they too may have some role here. Keep guessing and do contribute to the theory if you can!!
Now, I have discovered an article that links Glutamate to classical conditioning. It is titled Reward-Predictive Cues Enhance Excitatory Synaptic Strength onto Midbrain Dopamine Neurons, and here is the abstract:
Using sensory information for the prediction of future events is essential for survival. Midbrain dopamine neurons are activated by environmental cues that predict rewards, but the cellular mechanisms that underlie this phenomenon remain elusive. We used in vivo voltammetry and in vitro patch-clamp electrophysiology to show that both dopamine release to reward predictive cues and enhanced synaptic strength onto dopamine neurons develop over the course of cue-reward learning. Increased synaptic strength was not observed after stable behavioral responding. Thus, enhanced synaptic strength onto dopamine neurons may act to facilitate the transformation of neutral environmental stimuli to salient reward-predictive cues.
Though the article itself does not talk about glutamate, and nor does this Scicurious article on Neurotopia, commenting on the same , which focuses more on the dopamine connection, still I believe that we have a Glutamate connection here. First let us see how the artifact under discussion is indeed nothing but classical conditioning:
The basic idea is that, when you get a reward unexpectedly, you get a big spike of DA to make your brain go “sweet!” After a while, you being to recognize the cues behind the reward, and so seeing the wrapper to the candy will make your DA spike in anticipation. But it’s only very recently that we’ve been able to see this change taking place, and there were still lots of questions as to what was happening when these changes happen.
So the authors of this study took a bunch of rats. They implanted fast scan cyclic voltammetry probes into their heads. Voltammetry is a technique that allows you to detect changes in DA levels in brain areas (in this case the nucleus accumbens, an area linked with reward) which represent groups of cells firing. So the rats had probes in their heads detecting their DA, and then they were given a stimulus light (a conditioned stimulus), a nosepoke device, and a sugar pellet. There is nothing that a rat likes more than a sugar pellet, and so there was a nice big spike in DA as it got its reward. So the rats figured out pretty quickly that, when the light came on, you stick your nose in the hole, and sugar was on the way. As they learned the conditioned stimulus, their DA spikes in response to reward SHIFTED, moving backward in time, so that they soon got a spike of DA when they saw the light, without a spike when they got the pellet. This means that the animals had learned to associate a conditioned stimulus with reward. Not only that, the DA spike was higher immediately after learning than the spike in rats who just got rewards without learning.
So, if we consider the dopamine spike as an Unconditioned Response, then what we have is a new CS-> CR pairing or classical conditioning taking place. Now, the crucial study that showed that the learning is mediated by Glutamate: (emphasis mine)
To find out whether or not excitatory synapses were in fact changing, they authors conducted electrophysiology experiments in rats that were either trained or not trained. Electrophysiology is a technique where you actually put a tiny, tiny electrode into a cell membrane. When that cell is then stimulated, you can actually WATCH it fire. It’s really very cool to see. Of course all sorts of things are responsible for when a cell fires and how, but what they were looking at here were specific glutamate receptors known as AMPA and NMDA. These are two major receptors that receive glutamate currents, which are excitatory and induce cells downstream to fire. What they found was that, in animals that had been trained to a conditioned stimulus, AMPA and NMDA receptors had a much stronger influence on firing than in non-trained animals, which means that the synaptic strength on DA neurons is getting stronger as animals learn. Not only that, but cells from trained rats already exhibited long-term potentiation, a phenomenon associated with formation of things like learning and memory.
But of course, you have to make sure that glutamate is really the neurotransmitter responsible, and not just a symptom of something else changing. So they ran more rats on voltammetry and trained, and this time put a glutamate antagonist into the brain. The found that a glutamate antagonist completely blocked not only the DA shift to a conditioned stimulus, but the learning itself.
From the above it is clear that Glutamate , and the LTP that it leads to in the mid-brain neurons synapses , is crucial for Classical conditioning learning. Seems that one more puzzle is solved and another jig-jaw piece fits where it should have.