NONPARAMETRIC MODELS FOR DISCRIMINATION METHODS AND SENSITIVITY ANALYSIS FOR TRIADS

Nonparametric models for sensory discrimination methods are developed. In these models, the relationship between the probability of a correct response, P_c, and the measures for sensory discriminability or sensory difference, p₁ and p₂ are given. On the basis of the models, different sensitivities for triadic tests were compared including 3-AFC and triangle methods with the strong stimulus as the odd sample and with the weak stimulus as the odd sample. Predictions were made which can be tested experimentally.     

Optimum probabilistic processing in colour perception. II. Colour vision as template matching.

A statistical approach to account for psychophysical phenomena in human colour vision is presented. The central visual processor is viewed as an optimum recognizer of stochastic patterns supplied by the periphery. The processor makes an optimum estimate of the spectral parameters of the stimulus, given the wavelength filter characteristics of the periphery, the stochastic nature of the information and an internal template to which the external stimulus is matched. The estimate is constrained in ways inferred from empirical phenomena. Subjective brightness of monochromatic stimuli and related constant brightness manifolds in the colour space constitute the constraint for brightness estimation. Results analogous and in accord with those of earlier line element theories are obtained. The Bezold-Brücke hue shift constitutes the basic constraint for hue estimation. The hue estimate involves interrelation between the fields in the experiment. Similarities and differences both in basic conceptions and results introduced by the template matching notions are discussed.     

Multidimensional auditory stimulus control in a chimpanzee (Pan troglodytes)

An adolescent female chimpanzee (Pan troglodytes) was trained to discriminate auditory compound stimuli differing in tonal frequency and/or tone on-off rate. Following acquisition training and overtraining, she was shifted to multidimensional stimulus control testing using redundant relevant auditory stimulus sets with discriminability of elements in each dimension varied systematically. Although the control by both dimensions changed significantly as a function of discriminability, the degree of dimensional control was stronger in the tone on-off rate than in the tonal frequency. These results clearly demonstrated “attentional” control of the chimpanzee's auditory discrimination behavior and the interaction between two dimensions of auditory stimuli. The author is now at the Department of Psychology, Primate Research Institute, Kyoto University as a transfer student of the Doctor course.     

A random-walk interpretation of incentive effects in visual discrimination

Previous studies have shown that changes in stimulus discriminability and changes in reward density affect pigeon reaction-time (RT) distributions in different ways. A random-walk model (“RWP”) accounts for these differences and assigns a single parameter to each of the independent variables. This paper briefly reviews the model and illustrates its findings with hue discrimination data. A new analysis then presents fits to data showing that increased reward for stimulus “A” lengthens RT of pecks to an alternative stimulus “B”, and that this effect on RT distributions is much the same as the effect caused by reduction of reward to B. RWP account for both effects by changes in its “bias” parameter. The remainder of the paper comments on the relations between reward, RT, incentive and bias.     

Contextual conditioning. A comparison of eastern and western views

Eastern and Western interpretations of contextual control of phasic conditional responses (transswitching) are contrasted. The Eastern (Asratyan, 1965) approach emphasizes the role of the tonic conditional stimulus and the (hypothetical) tonic response it evokes. The Western (Lachnit, 1986) approach emphasizes the role of compound conditional stimuli. Although Lachnit showed that transswitching-like results can be obtained without a tonic stimulus, attempts to simulate transswitching experiments using a computer model of the Rescorla-Wagner theory (Kimmel and Lachnit, 1988) have shown that predictions from the theory approximate empirical results in human classical conditioning only when the tonic stimulus is given far greater weight than the phasic stimulus. In other words, only when the Rescorla-Wagner theory is made more like Asratyan's theory, can the compound conditional stimulus approach account for real empirical transswitching data.     

Classical conditioning,signal detection,and evolution

Strength of classical conditioning is increased either by increasing discriminability of the conditioned stimulus (CS) from the background, or by increasing contingency between conditioned and unconditioned stimili (US).Classical conditioning can be regarded as a decision process in which the subject has to decide whether or not to respond with a conditioned response in the presence or absence of the CS. According to modern evolutionary theories, it might be assumed that this decision process maximizes the trade-off between cost and benefits.By assuming that the decision rule maximizes expected benefit, the empirical relationship between contingency and the strength of classical conditioning is theoretically derived. In addition, when the decision rule is incorporated to a signal detection paradigm, theoretical results describing the relationship between CS discriminability and CS – US contingency with the strength of classical conditioning are in agreement with experimental data.     

The effect of stimulus number on the stability of responses for an extensive heat pain test

Background.?The use of relatively lower stimulus presentation numbers in quantitative sensory testing may influence the computation accuracy of participants’ discriminability. The minimum trial number for obtaining a stabilized participant discrimination ability was determined.

Materials and methods.?Twelve participants’ ability to discriminate between noxious heat stimuli pairs (45°C/46°C, 46°C/47°C, and 47°C/48°C) was assessed using a six-category confidence rating scale. Heat stimuli were administered to the forearm. Two conditions with presentation numbers of 17 trials per stimulus (representing the median number of trials in previous studies) and 40 trials per stimulus (used in a previous study with a similar protocol) were used.

Results and discussion.?Participants’ discriminability stabilized at approximately the 20th trial based on the lowest frequency of indeterminate and non-model conforming results under both conditions. A simple linear regression model showed a statistically significant positive relationship between discriminability for the two conditions (slope?=?0.65, p?<?0.001; constant?=?0.33, p?=?0.02; r^2?=?0.51). As a rule of thumb, approximately 20 trials per stimulus intensity could be used to obtain a stabilized discriminability outcome.     

Need probability effects in animal short-term memory

Five pigeons performed in a delayed matching-to-sample (DMTS) procedure with five delay durations (0.5, 2.5, 5, 10 and 20 s) mixed within sessions. Contrary to the predictions of need probability theory, discriminability decreased when fewer short than long delays were included in each session. To test whether the decrease in discriminability was due to a decrease in obtained reinforcement at short delays, the number of trials at each delay was held constant and reinforcer probability was increased with increasing delay. This manipulation produced a similar decrease in discriminability as when the frequency of delays was manipulated. It was concluded that the effect of delay frequency on the forgetting function is mediated by the effect of the reinforcer distribution, which influences discriminability by weakening stimulus control.     

Stimulus-dependent adjustment of reward prediction error in the midbrain

Previous reports have described that neural activities in midbrain dopamine areas are sensitive to unexpected reward delivery and omission. These activities are correlated with reward prediction error in reinforcement learning models, the difference between predicted reward values and the obtained reward outcome. These findings suggest that the reward prediction error signal in the brain updates reward prediction through stimulus-reward experiences. It remains unknown, however, how sensory processing of reward-predicting stimuli contributes to the computation of reward prediction error. To elucidate this issue, we examined the relation between stimulus discriminability of the reward-predicting stimuli and the reward prediction error signal in the brain using functional magnetic resonance imaging (fMRI). Before main experiments, subjects learned an association between the orientation of a perceptually salient (high-contrast) Gabor patch and a juice reward. The subjects were then presented with lower-contrast Gabor patch stimuli to predict a reward. We calculated the correlation between fMRI signals and reward prediction error in two reinforcement learning models: a model including the modulation of reward prediction by stimulus discriminability and a model excluding this modulation. Results showed that fMRI signals in the midbrain are more highly correlated with reward prediction error in the model that includes stimulus discriminability than in the model that excludes stimulus discriminability. No regions showed higher correlation with the model that excludes stimulus discriminability. Moreover, results show that the difference in correlation between the two models was significant from the first session of the experiment, suggesting that the reward computation in the midbrain was modulated based on stimulus discriminability before learning a new contingency between perceptually ambiguous stimuli and a reward. These results suggest that the human reward system can incorporate the level of the stimulus discriminability flexibly into reward computations by modulating previously acquired reward values for a typical stimulus.     

The behavioral specificity of stimulation: a systems approach to procedural distinctions of classical and instrumental conditioning.

This paper presents a new methodological approach to classical Pavlovian investigations. Using a yoked group for stimulus pairing trial pacing, behavioral occurrence frequencies, sequential behavior patterns, and behavioral change dynamics are analyzed for rats in both a non-discriminative and a general setting discrimination paradigm. Theoretical issues addressed by a variety of new dependent measures are discussed, and a special emphasis is placed on comparing instrumental and classical procedures as viewed from the behavioral perspective of the subject.     

Adaptive Spike Threshold Enables Robust and Temporally Precise Neuronal Encoding

Neural processing rests on the intracellular transformation of information as synaptic inputs are translated into action potentials. This transformation is governed by the spike threshold, which depends on the history of the membrane potential on many temporal scales. While the adaptation of the threshold after spiking activity has been addressed before both theoretically and experimentally, it has only recently been demonstrated that the subthreshold membrane state also influences the effective spike threshold. The consequences for neural computation are not well understood yet. We address this question here using neural simulations and whole cell intracellular recordings in combination with information theoretic analysis. We show that an adaptive spike threshold leads to better stimulus discrimination for tight input correlations than would be achieved otherwise, independent from whether the stimulus is encoded in the rate or pattern of action potentials. The time scales of input selectivity are jointly governed by membrane and threshold dynamics. Encoding information using adaptive thresholds further ensures robust information transmission across cortical states i.e. decoding from different states is less state dependent in the adaptive threshold case, if the decoding is performed in reference to the timing of the population response. Results from in vitro neural recordings were consistent with simulations from adaptive threshold neurons. In summary, the adaptive spike threshold reduces information loss during intracellular information transfer, improves stimulus discriminability and ensures robust decoding across membrane states in a regime of highly correlated inputs, similar to those seen in sensory nuclei during the encoding of sensory information.     

The effects of repetitive contractions and ischemia on the ability to discriminate intramuscular sensation

In order to investigate whether repetitive, low-level, muscular contractions or ischemia affect the ability of subjects to discriminate electrical stimuli delivered to the forearm musculature, a total of 25 experiments was performed on a group of five healthy subjects utilizing signal detection methodology. EMG needle electrodes were inserted into the forearm extensor musculature and the discriminability of two different pairs of constant current electrical stimuli were measured for sequential blocks of 100 trials, before and after the experimental interventions. Repetitive muscular activity consisting of a 15-min typing intervention at a rate of 60-80 wpm using a numeric keypad led to a significant decrease in the ability to discriminate the non-noxious stimulus pair in the block of trials immediately following the typing intervention, which then returned to pre-intervention discrimination levels following a 5-min break. The forearm ischemia significantly impaired the ability to discriminate both the noxious and non-noxious stimulus pair in the blocks of trials during and immediately after ischemia, which then returned to pre-intervention discrimination levels following a 5-min break. These experiments demonstrate that both repetitive muscular activity and ischemia acutely decrease the ability to discriminate intramuscular sensation. The mechanism may be due to decreased cortical processing, spinal cord sensitization or peripheral ischemia of large diameter afferents. These findings may be relevant to the physiological mechanisms underlying the development of overuse injuries.     

Biological probability: cognitive processes of generating probabilities of events in biological systems.

This paper analyses relationships between probabilities of events happening in biological systems (or probabilistic disposition of systems) and cognitive properties of biological entities comprising such systems. Two kinds of cognitive properties are identified as relevant to the current problem: the ability to respond differently against different configurations of the environment (discriminability of cognition), and the ability to make an appropriate response to maintain a particular relation with the environment (selectivity of cognition). A basic framework bridging the two features of living systems, probabilistic disposition and the cognitive properties, is presented towards a general theory explaining the process generating probabilities of biological events. In this framework, a deterministic model of a system of entities is developed, in which objects are described as subjects that cognize events (i.e. entities as cognizers). Cognition is used in a wider sense, including not only biotic but also abiotic, and cognizers are conceptually distinguished from the meta-observer who describes the system externally. Based on this perspective, this paper seeks to explicate how events can occur in an uncertain, probabilistic manner, if observed from a cognizer viewpoint, even under a deterministic system. Each cognizer is identified with both the set of states that are actually taken, and its motion function which maps its state uniquely to a successor state depending on the current states of itself and of the rest of cognizers constituting the system. The model analysis reveals that the cognitive properties, discriminability and selectivity, of a cognizer can contribute to determining the probability of an event encountered by the cognizer itself-in particular, discrimination reducing the uncertainty in events occurrence for the cognizer. Biological implication of this result is discussed focusing on the concept of the probability of survival and reproduction.     

Explaining the Timing of Natural Scene Understanding with a Computational Model of Perceptual Categorization

Observers can rapidly perform a variety of visual tasks such as categorizing a scene as open, as outdoor, or as a beach. Although we know that different tasks are typically associated with systematic differences in behavioral responses, to date, little is known about the underlying mechanisms. Here, we implemented a single integrated paradigm that links perceptual processes with categorization processes. Using a large image database of natural scenes, we trained machine-learning classifiers to derive quantitative measures of task-specific perceptual discriminability based on the distance between individual images and different categorization boundaries. We showed that the resulting discriminability measure accurately predicts variations in behavioral responses across categorization tasks and stimulus sets. We further used the model to design an experiment, which challenged previous interpretations of the so-called “superordinate advantage.” Overall, our study suggests that observed differences in behavioral responses across rapid categorization tasks reflect natural variations in perceptual discriminability.     

Reading spike timing without a clock: intrinsic decoding of spike trains

The precise timing of action potentials of sensory neurons relative to the time of stimulus presentation carries substantial sensory information that is lost or degraded when these responses are summed over longer time windows. However, it is unclear whether and how downstream networks can access information in precise time-varying neural responses. Here, we review approaches to test the hypothesis that the activity of neural populations provides the temporal reference frames needed to decode temporal spike patterns. These approaches are based on comparing the single-trial stimulus discriminability obtained from neural codes defined with respect to network-intrinsic reference frames to the discriminability obtained from codes defined relative to the experimenter''s computer clock. Application of this formalism to auditory, visual and somatosensory data shows that information carried by millisecond-scale spike times can be decoded robustly even with little or no independent external knowledge of stimulus time. In cortex, key components of such intrinsic temporal reference frames include dedicated neural populations that signal stimulus onset with reliable and precise latencies, and low-frequency oscillations that can serve as reference for partitioning extended neuronal responses into informative spike patterns.     

Neural Coding with Graded Membrane Potential Changes and Spikes

The neural encoding of sensory stimuli is usually investigated for spike responses, although many neurons are known to convey information by graded membrane potential changes. We compare by model simulations how well different dynamical stimuli can be discriminated on the basis of spiking or graded responses. Although a continuously varying membrane potential contains more information than binary spike trains, we find situations where different stimuli can be better discriminated on the basis of spike responses than on the basis of graded responses. Spikes can be superior to graded membrane potential fluctuations if spikes sharpen the temporal structure of neuronal responses by amplifying fast transients of the membrane potential. Such fast membrane potential changes can be induced deterministically by the stimulus or can be due to membrane potential noise that is influenced in its statistical properties by the stimulus. The graded response mode is superior for discrimination between stimuli on a fine time scale.     

How does the toad's visual system discriminate different worm-like stimuli?

Behavioral experiments show that toads exhibit stimulus- and locus-specific habituation. Different worm-like stimuli that toads can discriminate at a certain visual location form a dishabituation hierarchy. What is the neural mechanism which underlies these behaviors? This paper proposes that the toad discriminates visual objects based on temporal responses, and that discrimination is reflected in different average neuronal firing rates at some higher visual center, hypothetically anterior thalamus. This theory is developed through a large-scale neural simulation which includes retina, tectum and anterior thalamus. The neural model based on this theory predicts that retinal R2 cells play a primary role in the discrimination via tectal small pear cells (SP) and R3 cells refine the feature analysis by inhibition. The simulation demonstrates that the retinal response to the trailing edge of a stimulus is as crucial for pattern discrimination as the response to the leading edge. The new dishabituation hierarchies predicted by this model by reversing contrast and shrinking stimulus size need to be tested experimentally.     

A theory of natural selection incorporating interaction among individuals. I. The modeling process

Short- and long-term consequences of natural selection, operating in accordance with the classical (non-interaction) model, are reviewed. This review provides the basis for comparisons of the optimum results of the classical model with the results from interaction models. Then as a first step, the simplest interaction modeling system (Model I) is developed subject to the following conditions: (i) The natural selection model is constructed so that it is compatible with the one that already exists for artificial selection theory. This condition ensures that an overall theory is formulated which embraces both natural and artificial selection with a single modeling system and a common notation. (ii) The interaction theory is developed so that it is an extension of the classical theory. This condition ensures that the generalized theory includes the classical results as a special case when interaction is absent.The short- and long-term consequences of selection operating with Model I are determined to be less than optimal when compared with the classical results. Future papers in this series will consider how the interaction model can be modified in order to produce results which tend toward optimality.     

Absolute and relational control of a sequential auditory discrimination by pigeons (Columba livia)

Recent evidence indicates that pigeons can readily learn visual discriminations based on both absolute and relational stimulus factors. To examine how these two types of control function in their non-dominant auditory modality, we tested four pigeons in a go/no-go sequential auditory discrimination in which both absolute and relational cues were redundantly available. In this task, sequences of different sounds created from one set of pitches were reinforced, while different sequences created from another set of pitches and any same sequences made from either set of pitches were not. Across three experiments, we independently varied the relative discriminability of the absolute and relational components. The pigeons were consistently and primarily controlled by the absolute fundamental pitch of our notes in all of the experiments, although this was influenced by the range and arrangement of the pitches used in each set. A majority of the pigeons also demonstrated relational control when this component was made more salient. The more robust control exhibited by absolute factors is consistent with the comparative hypothesis that birds in general may have a well-developed aptitude for processing absolute pitch in many auditory settings. The relational control is consistent with our recent evidence of same/different auditory learning by pigeons.