Saturday, February 15, 2014

119. The Human Neocortex

The human brain, along with the spinal cord, comprises the central nervous system. The top outer portion of the brain, just under the scalp, is the neocortex (or cortex for short). It covers most of the R-brain (R for reptilian), and has a crumpled appearance, with many ridges and valleys. The R-brain is rather similar in reptiles and mammals, and has a number of parts, including the thalamus and the hippocampus.

Humans are special compared to other mammals because of their very prominent prefrontal cortex (or frontal lobe). The prefrontal cortex (particularly the upper two-thirds of it, including the dorsolateral prefrontal cortex) can be regarded as the rational centre of the brain; or the rational brain. The rest of it is the emotional brain.

The human cortex, if stretched flat, is the size of a large napkin, and ~2 mm thick. It has six layers, each roughly the thickness of a playing card. There is a branching hierarchy among the layers. Layer 6 is at the bottom of the hierarchy, and Layer1 is at the top. The inputs from the various sensory organs are received in Layer 6, and then interpreted and correlated. Then more and more abstract and generalized versions of the information are sent up the hierarchical layers. There is a very high degree of feedback and feedforward among the layers, as also cross-correlations.

There are ~1011 nerve cells or neurons in the human cortex. Most of them have a pyramidal-shaped central body or nucleus, as well as an axon, and a number of branching structures called dendrites. We can think of the axon as a signal emitter, and the dendrites as signal receivers. When a strand of an axon of one neuron (the presynaptic neuron) ‘touches’ a dendrite of another neuron (the postsynaptic neuron), a connection called a synapse is established. A typical axon is involved in several thousand synapses.

Portions of the cortex can be identified as different functional areas or regions. For example, a portion of the frontal lobe is the motor cortex. It controls movement and other actuator functions of the body.

The cortical tissue can be functionally divided into vertical units or columns. Neurons within a column respond in a similar manner to external signals with a particular attribute.

When a sensory or other pulse (‘spike’) involving a particular synapse arrives at the axon, it causes the synaptic vesicles in the presynaptic neuron to release chemicals called neurotransmitters into the gap or synaptic cleft between the axon of the first neuron and the dendrite of the second. These chemicals bind to the receptors on the dendrite, triggering a brief local depolarization of the membrane of the postsynaptic cell. This is described as a firing of the synapse by the presynaptic neuron.

If a synapse is made to fire repeatedly at high frequency, it becomes more sensitive; i.e. subsequent signals make it undergo greater voltage swings or spikes. Building up of memories amounts to formation and strengthening of synapses.

The firing of neurons follows two general rules:

(1) Neurons which fire together wire together. Connections between neurons firing together in response to the same signal get strengthened.

(2) Winner-takes-all inhibition. When several neighbouring neurons respond to the same input signal, the strongest or the ‘winner’ neuron will inhibit the neighbours from responding to the same signal in future. This makes these neighbouring neurons free to respond to other types of input signals.

The functionality of the cortex is arranged in a branching hierarchy. The primary sensory regions constitute the lowest rung of the hierarchy (Layer 6). The sensory region for, say, vision (called V1) is different from that for hearing etc. V1 feeds information to higher layers called V2, V4 and IT, and to some other regions. The higher they are in the hierarchy, the more abstract they become. V2, V4 etc. are concerned with more specialized or abstract aspects of vision. The higher echelons of the functional region responsible for vision have the visual memories of all sorts of objects. Similarly for other sensory perceptions.

In the higher echelons are areas called association areas. They receive inputs from several functional regions. For example, signals from both vision and audition reach one such association area.
Although the primary sensory mechanism for, for example, vision is not the same as for hearing, what reaches the brain at higher levels of the hierarchy is qualitatively the same. The axons carry neural signals or spikes which are partly chemical and partly electrical, but their nature is independent of whether the primary input signal was visual or auditory or tactile. Finally they are just patterns.

Creation of short-term memory in the brain amounts to a stimulation of the relevant synapses, which is enough to temporarily strengthen or sensitize them to subsequent signals.

This strengthening of the synapses becomes permanent in the case of long-term memory. This involves the activation of genes in the nuclei of postsynaptic neurons, initiating the production of proteins in them. Thus learning requires the synthesis of proteins in the brain within minutes of the training. Otherwise the memory fades away.

Information meant to become the higher-level or generalized memory, called declarative memory, passes through the hippocampus, before reaching the cortex. The hippocampus is like the principal server on a computer network. It plays a crucial role in consolidating long-term memories and emotions by integrating information coming from sensory inputs with information already stored in the brain.