From Wevision, a redoubtable and rich site on the retina and vision: There will be more data in a subsequent message. I lost, but will find, some PDFs on minimum number of rods to register an observer’s sense of an event – 5 to 14according to an article segment below – as well as the temporal resolution of the conduction, which I think is important. There is an article in WebVision on that, but I can barely understand it.
What can be gleaned from the data, aside from the fact that they’re cool? An honest question.
Rods peak in density 18o or 5mm out from the center of the fovea in a ring around the fovea at 160,000 rods/mm2. (Fig. 5)
No rods in central 200 µm.
Average 80-100,000 rods/mm2
Rod acuity peak is at 5.2o or 1.5 mm from foveal center where there are 100,000 rods/mm2 (Mariani et al.,1984).
Fig. 5. Density plot of rods and cones on the horizontal meridian of the human retina (59 K jpeg image)
There is a peak of the rod photoreceptors in a ring around the fovea at about 4.5 mm or 18 degrees from the foveal pit. I believe that the number and separation of rods is important in the projections of light distribution.
Rods convey the ability to see at night, under conditions of very dim illumination. Animals with high densities of rods tend to be nocturnal, whereas those with mainly cones tend to be diurnal. The nature of dim light is important both to physicists and to biologists. In 1905 Einstein proposed that light propagated only in discrete irreducible packets or quanta. This explained the non-classical features of the ‘photoelectric effect’, a process by which light releases electrons from metal surfaces, described by Heinrich Hertz in 1887. Rods are so sensitive that they actually detect single quanta of light, much as do the most sensitive of physical instruments. In 1942 Selig Hecht argued that human rods must be capable of detecting individual light quanta because light flashes so dim that only 1 in 100 rods were likely to absorb a quantum were yet reliably seen by careful observors. A century after the original discovery of the photoelectric effect it has become possible to record directly the minute electrical voltages in rods induced by absorption of individual light quanta. An excellent example is shown in the suction electrode recordings of monkey rods by Schneeweis and Schnapf (1995) (Fig. 22). Each dot in the figure below represents delivery of a very dim pulse of light containing only a few quanta. Voltage responses appear to come in 3 sizes: none, small, and large, representing the detection of 0, 1 or 2 quanta in each flash. The granularity of response to dim light stimuli is evident.
Fig. 22. Photovoltages recorded in monkey rods
Rod sensitivity appears to be bought at a price, however, since rods are much slower to respond to light stimulation than cones. This is one reason why sporting events such as baseball become progressively more difficult as daylight fails. Both electrical recordings and human observations suggest that signals from rods may arrive as much as 1/10 second later than those from cones under lighting conditions where both can be simultaneously activated (MacLeod, 1972).
- Can we see a single photon?
The minimum number of photons required to produce a visual effect was first successfully determined by Hecht, Schlaer and Pirenne in a landmark experiment (Hecht et al., 1942). Human subjects were allowed to stay in the dark for 30 minutes to have optimal visual sensitivity. The stimulus was presented 20 degrees to the left of the point of focus so that the light would fall on the region of the retina with the highest concentration of rods. The stimulus was a circle of red light with a diameter of 10 minutes (1 minute=1/60th of a degree). The subjects were asked whether they had seen a flash. The light was gradually reduced in intensity until the subjects could only guess the answer. It was found that between 54 and 148 photons were required in order to elicit visual experience. After corrections for corneal reflection (4%), ocular media absorption (50%) and photons passed through retina (80%), only 5 to 14 photons were actually absorbed by the retinal rods. The small number of photons in comparison with the large number of rods (500) involved makes it very unlikely that any rod will take up more than one photon. Therefore, one photon must be absorbed by each of 5 to 14 rods in the retina to produce a visual effect.
In the same publication (Hecht et al., 1942), Hecht and co-workers determined the visual threshold of human vision by the famous “frequency-of-seeing-curves” experiment. The theory is that photon absorptions vary according to a Poisson probability distribution. If a is the average number of photons absorbed per flash, the probability Pn that any number n will be absorbed is: Pn = an/(ean!). Some of these Possson integral curves (n from 1 to 9) were shown in Figure 5A. By measuring the frequency of seeing against the logarithm of the brightness experimentally, one can fit the experimental curve with one of the probability distribution in Figure 4A to reveal the value n, which lies between 5 and 8 (Figure 4B). This agrees well with the value 5 to 14 photons absorbed by rods!
Figure 4. Frequency of seeing curve experiment. (A) Poisson probability distribution. For any average number of quanta (hv) per flash, the ordinates give the probabilities that the flash will deliver to the retina n or more photons, depending on the value assumed for n. (B) Relation between the average energy content of a flash of light (in number of (hv) and the frequency with which it is seen by three observers. From Figs. 6 & 7 of (Hecht et al., 1942)