Curriculum Vitae - March 1995 Lance William HahnDepartment of Neuroscience 1007 S. 49th, 3rd Floor
123 Anatomy/Chemistry Bldg Philadelphia, PA 19143
University of Pennsylvania (215) 724-3435
Philadelphia, PA 19104-6058 (215) 898-7536
e-mail: lance@retina.anatomy.upenn.edu
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My broad research goal is to explain the way in which the human visual system transforms the images projected onto the two retinas into a cohesive visual percept. Two fundamental aspects of this transformation compose my main interest: spatial vision and adaptation. By understanding spatial vision one learns how information is derived from changes in light intensity across the retinal image. By understanding adaptation one learns how the visual system alters itself to become well-suited for the immediate environment. Rather than focusing solely on spatial vision or adaptation, my research examines the way in which adaptation changes spatial vision.
Adaptation is typically described as the way in which exposure to an adapting stimulus changes the perception of a target stimulus. In order to create a computational model of an adaptation process, at least two things need to be known about the adaptation mechanism: first, the nature of the signal producing the adaptation (for example, a light signal produces light adaptation) and second, a quantitative description of the way in which the adapting stimulus changes the perception of the target stimulus (for example, a reduction in the apparent intensity of a target signal by a multiplicative factor). This approach has been used to describe several light adaptation mechanisms and contrast adaptation mechanisms. I have used elaborations of this paradigm to distinguish between adaptation mechanisms and to make precise predictions about the way in which adaptation alters spatial vision in the dark and in an illuminated environment.
To test this hypothesis I created a five phase plan: 1) collect data describing the time-course of local light adaptation, 2) collect data describing eye movements during steady fixation, 3) form an integrated model of both phenomena, 4) use the model to make explicit predictions about the way in which the interaction between light adaptation and eye movements would change the contrast sensitivity function and 5) test the predictions empirically. I developed a model which was completely defined by measurements of adaptation and eye-movements when the phenomena were independent, but I used the model to predict contrast sensitivity when the two phenomena interacted. The change in the contrast sensitivity function predicted by the model fit the measured change quite well. Thus, it is very likely that the decrease in sensitivity to low spatial frequency gratings with long durations is due to an interaction between local light adaptation and eye-movements during steady fixation. Although there certainly is a center-surround antagonism in the retina, it is not responsible for this decrease in sensitivity to low spatial frequencies. Because the contrast sensitivity function is widely used in describing the human visual system and in implementing computer-based visual systems, I believe that this new understanding of sensitivity to low spatial frequency stimuli will have a wide-spread impact on the field.
Another goal of my dissertation was to measure the cumulative effect of adaptation mechanisms on spatial vision. As mentioned earlier, one way of categorizing adaptation mechanisms is by the type of signal which produces the adaptation (i.e. light or contrast). However, a distinction can also be made between mechanisms which are quantitatively different, even though they may act upon the same signal. Two classes of adaptation mechanism which are characterized by their quantitative descriptions are subtractive adaptation, in which a portion of the adapting signal is subtracted from the target signal, and multiplicative adaptation, in which the target signal is divided by a portion of the adapting signal. These distinctions result in a set of four well-defined adaptation mechanisms: subtractive light adaptation, subtractive contrast adaptation, multiplicative light adaptation and multiplicative contrast adaptation.
Typically these four adaptation mechanisms are studied in isolation, but in the natural environment, these mechanisms almost certainly act in concert. One goal of my dissertation was to measure the coexisting influence of these mechanisms on spatial vision. I used a contrast matching task which allowed me to simultaneously determine the relative contributions of three of the four types of adaptation. After considering many plausible combinations of the four adaptation mechanisms, I concluded that the results were best described by a model which included only the multiplicative light adaptation mechanism. Although the simple model provided a good fit to the data, there were systematic differences between the data and the model which suggest other mechanism(s) are at work. Developing hypotheses about the missing component(s) in the model will be a goal of future research. Mechanisms which I plan pursue as possible components will include: luminance nonlinearities, cortical contrast nonlinearities and the division of stimulus parameters between different neural channels. All of these components can be expressed within the framework of a computational model.
In the work I have described so far the adapting stimulus has been a pattern - a sinewave grating. A sinewave grating can produce both light adaptation and contrast adaptation. Isolated light adaptation can be produced by changing the mean light level. This is the adaptation one experiences when leaving a dark movie theater for a sunlit parking lot. Classic measurements of the change in spatial vision produced by changes in the state of light adaptation have been made by measuring contrast sensitivity functions for different levels of background light. While the change in spatial vision as a function of background light level is well-documented, the change in spatial vision during dark adaptation is surprisingly undocumented. Furthermore, two existing hypotheses (a local multiplicative light adaptation hypothesis and an equivalent background hypothesis) make conflicting predictions about how spatial vision changes during dark adaptation. So, in a collaboration with Bill Geisler which preceded my dissertation (Hahn & Geisler, 1995), I resolved to measure contrast sensitivity as a function of time following exposure to an intense adapting field.
We traced the change in sensitivity of observers to a new spatial stimulus following exposure to a uniform field of intense light. (It was necessary to develop a new type of stimulus to make the measurements in the dark.) We also measured changes in the detectability of the new stimulus for different background light levels. This latter measurement allowed us to compare our measurements with other classic measurements to insure that the new stimulus was functionally similar to the traditional stimuli.
Our results showed that long-term dark adaptation produces the same increase in sensitivity at low spatial frequencies (coarse gratings) as it produces at high spatial frequencies (fine gratings). This finding supports the local multiplicative luminance adaptation hypothesis, and rejects the equivalent background hypothesis. In addition to discriminating between these two hypotheses, the data provide a singular description of how spatial vision changes during long-term dark adaptation.
Finally, I also plan to test the limits of existing models of adaptation and spatial vision with more complex stimuli. Historically the literature has been dominated by a motivation to isolate and measure a single attribute of adaptation or spatial vision. This has meant using mathematically simple stimuli which are very artificial. I will test the models which predict response to simple stimuli with stimulus configurations which approach the complexity of stimuli found in the natural environment. The use of more complex stimuli is likely to introduce new types of adaptation mechanism, new types of spatial mechanisms, and new space-adaptation interactions. For instance, it is known that prolonged exposure to a drifting grating produces motion adaptation. Although some spatial attributes of motion adaptation are known, little is known about interactions between motion adaptation other types of adaptation. In cooperation with computational modeling, the study of visual sensation has produced a collection of functional building blocks which I intend to use to form more general models of visual perception.