Abstract: Classically, biologists have considered adaptation of behavioural characteristics in terms of long-term functional benefits to the individual, such as survival or reproductive fitness. In captive species, including the domestic horse, this level of explanation is limited, as for the most part, horses are housed in conditions that differ markedly from those in which they evolved. In addition, an individual horse's reproductive fitness is largely determined by man rather than its own behavioural strategies. Perhaps for reasons of this kind, explanations of behavioural adaptation to environmental challenges by domestic animals, including the capacity to learn new responses to these challenges, tend to concentrate on the proximate causes of behaviour. However, understanding the original function of these adaptive responses can help us explain why animals perform apparently novel or functionless activities in certain housing conditions and may help us to appreciate what the animal welfare implications might be. This paper reviews the behavioural adaptation of the domestic horse to captivity and discusses how apparently abnormal behaviour may not only provide a useful practical indicator of specific environmental deficiencies but may also serve the animal as an adaptive response to these deficiencies in an “abnormal” environment.

Abstract: Feeding and training may affect acid-base responses to strenuous exercise. Acidosis usually correlates with higher blood lactate concentrations during intense exercise, but alkalosis has been found in several studies of horses, and higher lactate responses during sprints have been found in fat adapted horses. To elucidate these unexpected findings, we applied a comprehensive physicochemical approach to evaluate acid-base responses during exercise in fat adapted horses. In incremental tests and repeated sprints, changes in blood [H+] were dependent upon corresponding changes in pCO2 but not strong ion difference (SID, the algebraic sum of ions of sodium, potassium, chloride and lactate). The influence of changes in [Lac-] were largely offset by changes in [Na+], [K+] and [Cl-], so that SID was unchanged and did not contribute to the exercise induced acidemia, so it may be inaccurate to term this a lacticacidosis. During repeated sprints, central venous [H+] increased (acidosis) but arterial [H+] decreased (alkalosis). These changes were consistent with concurrent changes in venous and arterial pCO2 but not SID. Fat adaptation decreased mixed venous pCO2 during repeated sprints, which is consistent with the lower respiratory quotient associated with fat oxidation. Less pulmonary work to eliminate CO2 could benefit horses under hot and humid conditions, especially those with mildly reduced pulmonary function. The blood lactate response was decreased during aerobic tests but increased during anaerobic tests on fat adapted horses. Fat adaptation appears to facilitate the metabolic regulation of glycolysis, by sparing glucose and glycogen at work of low intensity, but by promoting glycolysis when power is needed for high intensity exercise. The blood lactate response to repeated sprints was increased more by the combination of fat adaptation and oral supplementation of sodium bicarbonate than by the sum of the responses to fat alone or bicarbonate alone. This synergism suggests that need for further studies of the interaction of fat adaptation with dietary cation-anion balance, especially under hot conditions. These results integrate harmoniously with previous findings of lower feed intake and fecal output, lower loads of heat and CO2, lower water losses in the feces and by evaporation, and less spontaneous activity and reactivity in fat adapted horses. Thus fat adaptation confers several advantages on horses and presumably other equids used for hard work, especially in the heat.

Abstract: Objective The goal of this work is to study the dark adaptation curve of the normal horse electroretinogram (ERG). Procedures The electroretinographic responses were recorded from six healthy female ponies using a contact lens electrode and a mini-Ganzfeld electroretinographic unit. The horses were sedated intravenously with detomidine, an auriculopalpebral nerve block was then performed, and the pupil was fully dilated. The ERG was recorded in response to a low intensity light stimulus (30 mcd.s/m2) that was given at times (T) T = 5, 10, 15, 20, 25, 30, 40, 50, and 60 min of dark adaptation. Off-line analysis of the ERG was then performed. Results Mean b-wave amplitude of the full-field ERG increased continuously from 5 to 25 min of dark adaptation. The b-wave amplitude peaked at T = 25, however, there was no statistical significance between T = 20 and T = 25. The b-wave amplitude then remained elevated with no significant changes until the end of the study at T = 60 (P > 0.49). The b-wave implicit time increased continuously between T = 5 and T = 20, then gradually decreased until T = 60. No distinct a-wave was observed during the testing time. Conclusions Evaluation of horse rod function or combined rod/cone function by means of full-field ERG should be performed after a minimum 20 min of dark adaptation.

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: Socio-ecological explanations for intra- and interspecific variation in the social and spatial organization of animals predominate in the scientific literature. The socio-ecological model, developed first for the Bovidae and Cervidae, is commonly applied more widely to other groups including the Equidae. Intraspecific comparisons are particularly valuable because they allow the role of environment and demography on social and spatial organization to be understood while controlling for phylogeny or morphology which confound interspecific comparisons. Feral horse (Equus caballus Linnaeus 1758) populations with different demography inhabit a range of environments throughout the world. I use 56 reports to obtain 23 measures or characteristics of the behaviour and the social and spatial organization of 19 feral horse populations in which the environment, demography, management, research effort and sample size are also described. Comparison shows that different populations had remarkably similar social and spatial organization and that group sizes and composition, and home range sizes varied as much within as between populations. I assess the few exceptions to uniformity and conclude that they are due to the attributes of the studies themselves, particularly to poor definition of terms and inadequate empiricism, rather than to the environment or demography per se. Interspecific comparisons show that equid species adhere to their different social and spatial organizations despite similarities in their environments and even when species are sympatric. Furthermore, equid male territoriality has been ill-defined in previous studies, observations presented as evidence of territoriality are also found in non-territorial equids, and populations of supposedly territorial species demonstrate female defence polygyny. Thus, territoriality may not be a useful categorization in the Equidae. Moreover, although equid socio-ecologists have relied on the socio-ecological model derived from the extremely diverse Bovidae and Cervidae for explanations of variation in equine society, the homomorphic, but large and polygynous, and monogeneric Equidae do not support previous socio-ecological explanations for relationships between body size, mating system and sexual dimorphism in ungulates. Consequently, in spite of the efforts of numerous authors during the past two decades, functional explanations of apparent differences in feral horse and equid social and spatial organization and behaviour based on assumptions of their current utility in the environmental or demographic context remain unconvincing. Nevertheless, differences in social cohesion between species that are insensitive to intra- and interspecific variation in habitat and predation pressure warrant explanation. Thus, I propose alternative avenues of inquiry including testing for species-specific differences in inter-individual aggression and investigating the role of phylogenetic constraints in equine society. The Equidae are evidence of the relative importance of phylogeny and biological structure, and unimportance of the present-day environment, in animal behaviour and social and spatial organization.