Heffner, R. S., & Heffner, H. E. (1986). Localization of tones by horses: use of binaural cues and the role of the superior olivary complex. Behav Neurosci, 100(1), 93–103.
Abstract: The ability of horses to use binaural time and intensity difference cues to localize sound was assessed in free-field localization tests by using pure tones. The animals were required to discriminate the locus of a single tone pip ranging in frequency from 250 Hz to 25 kHz emitted by loudspeakers located 30 degrees to the left and right of the animals' midline (60 degrees total separation). Three animals were tested with a two-choice procedure; 2 additional animals were tested with a conditioned avoidance procedure. All 5 animals were able to localize 250 Hz, 500 Hz, and 1 kHz but were completely unable to localize 2 kHz and above. Because the frequency of ambiguity for the binaural phase cue delta phi for horses in this test was calculated to be 1.5 kHz, these results indicate that horses can use binaural time differences in the form of delta phi but are unable to use binaural intensity differences. This finding was supported by an unconditioned orientation test involving 4 additional horses, which showed that horses correctly orient to a 500-Hz tone pip but not to an 8-kHz tone pip. Analysis of the superior olivary complex, the brain stem nucleus at which binaural interactions first take place, reveals that the lateral superior olive (LSO) is relatively small in the horse and lacks the laminar arrangement of bipolar cells characteristic of the LSO of most mammals that can use binaural delta I.
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Friederici, A. D., & Alter, K. (2004). Lateralization of auditory language functions: a dynamic dual pathway model. Brain Lang, 89(2), 267–276.
Abstract: Spoken language comprehension requires the coordination of different subprocesses in time. After the initial acoustic analysis the system has to extract segmental information such as phonemes, syntactic elements and lexical-semantic elements as well as suprasegmental information such as accentuation and intonational phrases, i.e., prosody. According to the dynamic dual pathway model of auditory language comprehension syntactic and semantic information are primarily processed in a left hemispheric temporo-frontal pathway including separate circuits for syntactic and semantic information whereas sentence level prosody is processed in a right hemispheric temporo-frontal pathway. The relative lateralization of these functions occurs as a result of stimulus properties and processing demands. The observed interaction between syntactic and prosodic information during auditory sentence comprehension is attributed to dynamic interactions between the two hemispheres.
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Carroll, J., Murphy, C. J., Neitz, M., Hoeve, J. N., & Neitz, J. (2001). Photopigment basis for dichromatic color vision in the horse. J Vis, 1(2), 80–87.
Abstract: Horses, like other ungulates, are active in the day, at dusk, dawn, and night; and, they have eyes designed to have both high sensitivity for vision in dim light and good visual acuity under higher light levels (Walls, 1942). Typically, daytime activity is associated with the presence of multiple cone classes and color-vision capacity (Jacobs, 1993). Previous studies in other ungulates, such as pigs, goats, cows, sheep and deer, have shown that they have two spectrally different cone types, and hence, at least the photopigment basis for dichromatic color vision (Neitz & Jacobs, 1989; Jacobs, Deegan II, Neitz, Murphy, Miller, & Marchinton, 1994; Jacobs, Deegan II, & Neitz, 1998). Here, electroretinogram flicker photometry was used to measure the spectral sensitivities of the cones in the domestic horse (Equus caballus). Two distinct spectral mechanisms were identified and are consistent with the presence of a short-wavelength-sensitive (S) and a middle-to-long-wavelength-sensitive (M/L) cone. The spectral sensitivity of the S cone was estimated to have a peak of 428 nm, while the M/L cone had a peak of 539 nm. These two cone types would provide the basis for dichromatic color vision consistent with recent results from behavioral testing of horses (Macuda & Timney, 1999; Macuda & Timney, 2000; Timney & Macuda, 2001). The spectral peak of the M/L cone photopigment measured here, in vivo, is similar to that obtained when the gene was sequenced, cloned, and expressed in vitro (Yokoyama & Radlwimmer, 1999). Of the ungulates that have been studied to date, all have the photopigment basis for dichromatic color vision; however, they differ considerably from one another in the spectral tuning of their cone pigments. These differences may represent adaptations to the different visual requirements of different species.
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Wolfe, J. M. (1983). Hidden visual processes. Sci Am, 248(2), 94–103.
Abstract: Isoluminant stimulus is an image whose edges are defined only by a change in color, not by change in brightness. The stimulus here is imperfect: the blue parts and the green parts of the image are only as nearly equal in brightness as they can be on the printed page. Moreover, the change in brightness beyond the edge of the page is apparent, and so is the fact that the reader is holding the magazine at reading distance. When such cues are removed under laboratory conditions, subjects faced with an isoluminant stimulus prove unable to bring its edges into focus. This deficiency contributes to making a familiar face hard to recognize. The experiment indicates that the brain process underlying visual accommodation (the focusing of the eyes) cannot “see” color; it is a hidden process distinct from the processes that lead to perception. The image shows Groucho Marx as he appeared in the motion picture Horse Feathers.
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Timney, B., & Keil, K. (1999). Local and global stereopsis in the horse. Vision Res, 39(10), 1861–1867.
Abstract: Although horses have laterally-placed eyes, there is substantial binocular overlap, allowing for the possibility that these animals have stereopsis. In the first experiment of the present study we measured local stereopsis by obtaining monocular and binocular depth thresholds for renal depth stimuli. On all measures, the horses' binocular performance was superior to their monocular. When depth thresholds were obtained, binocular thresholds were several times superior to those obtained monocularly, suggesting that the animals could use stereoscopic information when it was available. The binocular thresholds averaged about 15 min arc. In the second experiment we obtained evidence for the presence of global stereopsis by testing the animals' ability to discriminate between random-dot stereograms with and without consistent disparity information. When presented with such stimuli they showed a strong preference for the cyclopean equivalent of the positive stimulus with the real depth. These results provide the first behavioral demonstration of a full range of stereoscopic skills in a lateral-eyed mammal.
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