Sensory Systems/Visual System/old/Signal Processing

As mentioned before the retina is the main component in the eye, because it contains all the light sensitive cells. Without it, the eye would be comparable to a digital camera without the CCD (Charge Coupled Device) sensor. This part elaborates on how the retina perceives the light, how the optical signal is transmitted to the brain and how the brain processes the signal to form enough information for decision making.

Creation of the initial signals - Photosensor Function

Vision invariably starts with light hitting the photo-sensitive cells found in the retina. Light-absorbing visual pigments, a variety of enzymes and transmitters in retinal rods and cones will initiate the conversion from visible EM stimuli into electrical impulses, in a process known as photoelectric transduction. Using rods as an example, the incoming visible EM hits rhodopsin molecules, transmembrane molecules found in the rods' outer disk structure. Each rhodopsin molecule consists of a cluster of helices called opsin that envelop and surround 11-cis retinal, which is the part of the molecule that will change due to the energy from the incoming photons. In biological molecules, moieties, or parts of molecules that will cause conformational changes due to this energy is sometimes referred to as chromophores. 11-cis retinal straightens in response to the incoming energy, turning into retinal (all-trans retinal), which forces the opsin helices further apart, causing particular reactive sites to be uncovered. This "activated" rhodopsin molecule is sometimes referred to as Metarhodopsin II. From this point on, even if the visible light stimulation stops, the reaction will continue. The Metarhodopsin II can then react with roughly 100 molecules of a G_s protein called transducing, which then results in α_s and βγ after the GDP is converted into GTP. The activated α_s-GTP then binds to cGMP-phosphodiesterase(PDE), suppressing normal ion-exchange functions, which results in a low cytosol concentration of cation ions, and therefore a change in the polarisation of the cell.

The natural photoelectric transduction reaction has an amazing power of amplification. One single retinal rhodopsin molecule activated by a single quantum of light causes the hydrolysis of up to 10⁶ cGMP molecules per second.

Photo Transduction

Representation of molecular steps in photoactivation (modified from Leskov et al., 2000). Depicted is an outer membrane disk in a rod. Step 1: Incident photon (hν) is absorbed and activates a rhodopsin by conformational change in the disk membrane to R*. Step 2: Next, R* makes repeated contacts with transducin molecules, catalyzing its activation to G* by the release of bound GDP in exchange for cytoplasmic GTP. The α and γ subunits Step 3: G* binds inhibitory γ subunits of the phosphodiesterase (PDE) activating its α and β subunits. Step 4: Activated PDE hydrolyzes cGMP. Step 5: Guanylyl cyclase (GC) synthesizes cGMP, the second messenger in the phototransduction cascade. Reduced levels of cytosolic cGMP cause cyclic nucleotide gated channels to close preventing further influx of Na+ and Ca2+.

1. A light photon interacts with the retinal in a photoreceptor. The retinal undergoes isomerisation, changing from the 11-cis to all-trans configuration.
2. Retinal no longer fits into the opsin binding site.
3. Opsin therefore undergoes a conformational change to metarhodopsin II.
4. Metarhodopsin II is unstable and splits, yielding opsin and all-trans retinal.
5. The opsin activates the regulatory protein transducin. This causes transducin to dissociate from its bound GDP, and bind GTP, then the alpha subunit of transducin dissociates from the beta and gamma subunits, with the GTP still bound to the alpha subunit.
6. The alpha subunit-GTP complex activates phosphodiesterase.
7. Phosphodiesterase breaks down cGMP to 5'-GMP. This lowers the concentration of cGMP and therefore the sodium channels close.
8. Closure of the sodium channels causes hyperpolarization of the cell due to the ongoing potassium current.
9. Hyperpolarization of the cell causes voltage-gated calcium channels to close.
10. As the calcium level in the photoreceptor cell drops, the amount of the neurotransmitter glutamate that is released by the cell also drops. This is because calcium is required for the glutamate-containing vesicles to fuse with cell membrane and release their contents.
11. A decrease in the amount of glutamate released by the photoreceptors causes depolarization of On center bipolar cells (rod and cone On bipolar cells) and hyperpolarization of cone Off bipolar cells.

Without visible EM stimulation, rod cells containing a cocktail of ions, proteins and other molecules, have membrane potential differences of around -40mV. Compared to other nerve cells, this is quite high (-65mV). In this state, the neurotransmitter glutamate is continuously released from the axon terminals and absorbed by the neighbouring bipolar cells. With incoming visble EM and the previously mentioned cascade reaction, the potential difference drops to -70mV. This hyper-polarisation of the cell causes a reduction in the amount of released glutamate, thereby affecting the activity of the bipolar cells, and subsequently the following steps in the visual pathway.

Similar processes exist in the cone-cells and in photosensitive ganglion cells, but make use of different opsins. Photopsin I through III (yellowish-green, green and blue-violet respectively) are found in the three different cone cells and melanopsin (blue) can be found in the photosensitive ganglion cells.

Processing Signals in the Retina

Different bipolar cells react differently to the changes in the released glutamate. The so called ON and OFF bipolar cells are used to form the direct signal flow from cones to bipolar cells. The ON bipolar cells will depolarise by visible EM stimulation and the corresponding ON ganglion cells will be activated. On the other hand the OFF bipolar cells are hyper polarised by the visible EM stimulation, and the OFF ganglion cells are inhibited. This is the basic pathway of the Direct signal flow. The Lateral signal flow will start from the rods, then go to the bipolar cells, the amacrine cells, and the OFF bipolar cells inhibited by the Rod-amacrine cells and the ON bipolar cells will stimulated via an electrical synapse, after all of the previous steps, the signal will arrive at the ON or OFF ganglion cells and the whole pathway of the Lateral signal flow is established.

When the action potential (AP) in ON, ganglion cells will be triggered by the visible EM stimulus. The AP frequency will increase when the sensor potential increases. In other words, AP depends on the amplitude of the sensor's potential. The region of ganglion cells where the stimulatory and inhibitory effects influence the AP frequency is called receptive field (RF). Around the ganglion cells, the RF is usually composed of two regions: the central zone and the ring-like peripheral zone. They are distinguishable during visible EM adaptation. A visible EM stimulation on the centric zone could lead to AP frequency increase and the stimulation on the periphery zone will decrease the AP frequency. When the light source is turned off the excitation occurs. So the name of ON field (central field ON) refers to this kind of region. Of course the RF of the OFF ganglion cells act the opposite way and is therefore called "OFF field" (central field OFF). The RFs are organised by the horizontal cells. The impulse on the periphery region will be impulsed and transmitted to the central region, and there the so-called stimulus contrast is formed. This function will make the dark seem darker and the light brighter. If the whole RF is exposed to light. the impulse of the central region will predominate.

Signal Transmission to the Cortex

As mentioned previously, axons of the ganglion cells converge at the optic disk of the retina, forming the optic nerve. These fibres are positioned inside the bundle in a specific order. Fibres from the macular zone of the retina are in the central portion, and those from the temporal half of the retina take up the periphery part. A partial decussation or crossing occurs when these fibres are outside the eye cavity. The fibres from the nasal halves of each retina cross to the opposite halves and extend to the brain. Those from the temporal halves remain uncrossed. This partial crossover is called the optic chiasma, and the optic nerves past this point are called optic tracts, mainly to distinguish them from single-retinal nerves. The function of the partial crossover is to transmit the right-hand visual field produced by both eyes to the left-hand half of the brain only and vice versa. Therefore the information from the right half of the body, and the right visual field, is all transmitted to the left-hand part of the brain when reaches the posterior part of the fore-brain (diencephalon).

The pathway to the central cortex

The information relay between the fibers of optic tracts and the nerve cells occurs in the lateral geniculate bodies, the central part of the visual signal processing, located in the thalamus of the brain. From here the information is passed to the nerve cells in the occipital cortex of the corresponding side of the brain. Connections from the retina to the brain can be separated into a 'parvocellular pathway' and a "magnocellular pathway". The parvocellular pathways signals color and fine detail, whereas the magnocellular pathways detect fast moving stimuli.

Connections from the retina to the brain can be separated into a "parvocellular pathway" and a "magnocellular pathway". The parvocellular pathway originates in midget cells in the retina, and signals color and fine detail; magnocellular pathway starts with parasol cells, and detects fast moving stimuli.

Signals from standard digital cameras correspond approximately to those of the parvocellular pathway. To simulate the responses of parvocellular pathways, researchers have been developing neuromorphic sensory systems, which try to mimic spike-based computation in neural systems. Thereby they use a scheme called "address-event representation" for the signal transmission in the neuromorphic electronic systems (Liu and Delbruck 2010 [1]).

Anatomically, the retinal Magno and Parvo ganglion cells respectively project to 2 ventral magnocellular layers and 4 dorsal parvocellular layers of the Lateral Geniculate Nucleus (LGN). Each of the six LGN layers receives inputs from either the ipsilateral or contralateral eye, i.e., the ganglion cells of the left eye cross over and project to layer 1, 4 and 6 of the right LGN, and the right eye ganglion cells project (uncrossed) to its layer 2, 3 and 5. From here the information from the right and left eye is separated.

Although human vision is combined by two halves of the retina and the signal is processed by the opposite cerebral hemispheres, the visual field is considered as a smooth and complete unit. Hence the two visual cortical areas are thought of as being intimately connected. This connection, called corpus callosum is made of neurons, axons and dendrites. Because the dendrites make synaptic connections to the related points of the hemispheres, electric simulation of every point on one hemisphere indicates simulation of the interconnected point on the other hemisphere. The only exception to this rule is the primary visual cortex.

The synapses are made by the optic tract in the respective layers of the lateral geniculate body. Then these axons of these third-order nerve cells are passed up to the calcarine fissure in each occipital lobe of the cerebral cortex. Because bands of the white fibres and axons pair from the nerve cells in the retina go through it, it is called the striate cortex, which incidentally is our primary visual cortex, sometimes known as V1. At this point, impulses from the separate eyes converge to common cortical neurons, which then enables complete input from both eyes in one region to be used for perception and comprehension. Pattern recognition is a very important function of this particular part of the brain, with lesions causing problems with visual recognition or blindsight.

Based on the ordered manner in which the optic tract fibres pass information to the lateral geniculate bodies and after that pass in to the striate area, if one single point stimulation on the retina was found, the response which produced electrically in both lateral geniculate body and the striate cortex will be found at a small region on the particular retinal spot. This is an obvious point-to-point way of signal processing. And if the whole retina is stimulated, the responses will occur on both lateral geniculate bodies and the striate cortex gray matter area. It is possible to map this brain region to the retinal fields, or more usually the visual fields.

Any further steps in this pathway is beyond the scope of this book. Rest assured that, many further levels and centres exist, focusing on particular specific tasks, like for example colour, orientations, spatial frequencies, emotions etc.

Visual Perception

Equipped with a firmer understanding of some of the more important concepts of the signal processing in the visual system, comprehension or perception of the processed sensory information is the last important piece in the puzzle. Visual perception is the process of translating information received by the eyes into an understanding of the external state of things. It makes us aware of the world around us and allows us to understand it better. Based on visual perception we learn patterns which we then apply later in life and we make decisions based on this and the obtained information. In other words, our survival depends on perception.