Abstract: An attempt to replicate Grzimek (1952; Z. Tierpsychol., 27: 330-338) is reported where a Quarter-Horse mare chose between colored and gray stimuli for food reinforcement. Stimuli varied across a broad range of reflectance values. A double-blind procedure with additional controls for auditory, olfactory, tactile, and position cues was used. The subject could reliably discriminate blue (462 nm) vs. gray, and red (700 nm) vs. gray without regard to reflectance (P<0.001), but could not discriminate green (496 nm) vs. gray. It is suggested that horses are dichromats in a manner similar to swine and cattle.

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.

Abstract: In the past, equine color vision was tested with stimuli composed either of painted cards or photographic slides or through physiological testing using electroretinogram flicker photometry. Some studies produced similar results, but others did not, demonstrating that there was not yet a definitive answer regarding color vision in horses (Equus caballus). In this study, a pseudoisochromatic plate test--which is highly effective in testing color vision both in small children and in adult humans--was used for the first time on a nonhuman animal. Stimuli consisted of different colored dotted circles set against backgrounds of varying dots. The coloration of the circles corresponded to the visual capabilities of different types of color deficiencies (anomalous trichromacy and dichromacy). Four horses were tested on a 2-choice discrimination task. All horses successfully reached criterion for gray circles and demonstration circles. None of the horses were able to discriminate the protan-deutan plate or the individual protan or deutan plates. However, all were able to discriminate the tritan plate. The results suggest that horses are dichromats with color vision capabilities similar to those of humans with red-green color deficiencies.

Abstract: This study replicated an unreported finding observed in a color perception experiment (Pick, Lovell, Brown, & Dail, 1994) where, after using the method of successive approximations to train a blue-gray discrimination, red-gray trials were initiated without further training. Although a gray choice had never been reinforced, the subject chose gray on the first 20 trials (p < .000001). In the study reported here, a horse was trained to approach a red feed bucket and not a green feed bucket. After the subject mastered the discrimination, a blue bucket was substituted for the previously reinforced red bucket. With double-blind controls in place, the subject chose the unreinforced green bucket on 15 out of the first 20 blue-green trials yielding a binomial p = 0.0148 that this outcome could be due to chance alone. These results are contrary to all behavioristic psychological learning theories, but consistent with prospect theory (Kahneman & Tversky, 1979). Prospect theory predicts that given a choice between two previously unreinforced stimuli, one familiar and the other novel, humans will choose the familiar. It is argued that the bias toward the familiar is the basis to a heuristic that has a genetic origin and should exist in other animals on the phylogenetic scale. The results of this study indicate that the heuristic is available at least as far down the scale as the horse. Conceptual replications using shape stimuli and sound stimuli are in progress.

Abstract: Six horses (Equus caballus) were trained to discriminate color from grays in a counterbalanced sequence in which lightness cues were irrelevant. Subsequently, the pretrained colors were presented in a different sequence. Two sets of novel colors paired with novel grays were also tested. Performance was just as good in these transfer tests. Once the horse had learned to select the chromatic from the achromatic stimulus, regardless of the specific color, they were immediately able to apply this rule to novel stimuli. In terms of the underlying visual mechanisms, the present study showed for the first time that the spectral sensitivity of horse cone photopigments, measured as cone excitation ratios, was correlated with color discrimination performance, measured as accuracy, repeated errors, and latency of approach.