Learning tests in animal psychology are highly standardized with regard to both the formulation of scientific questions and the methods employed to obtain reliable answers. The Skinnerian technique of conditioning rats or pigeons in small test chambers has established itself as the main laboratory method in this field. Despite the undeniable advantages of highly controlled and manipulated test conditions, a number of problems with respect to the welfare of the test animals arise from the standard application of this method. These problems are specified here along with the presentation of new techniques that have been developed at the University of Vienna. The methodological improvements are related to the housing, testing and weighing of the pigeons and ameliorate the conditions these animals previously confronted within the laboratory.

Behaving organisms are continually choosing. Recently the theoretical and empirical study of decision making by behavioral ecologists and experimental psychologists have converged in the area of foraging, particularly food acquisition. This convergence has raised the interdisciplinary question of whether principles that have emerged from the study of decision making in the operant conditioning laboratory are consistent with decision making in naturally occurring foraging. One such principle, the “parameter-free delay-reduction hypothesis, ” developed in studies of choice in the operant conditioning laboratory, states that the effectiveness of a stimulus as a reinforcer may be predicted most accurately by calculating the decrease in time to food presentation correlated with the onset of the stimulus, relative to the length of time to food presentation measured from the onset of the preceding stimulus. Since foraging involves choice, the delay-reduction hypothesis may be extended to predict aspects of foraging. We discuss the strategy of assessing parameters of foraging with operant laboratory analogues to foraging. We then compare the predictions of the delay-reduction hypothesis with those of optimal foraging theory, developed by behavioral ecologists, showing that, with two exceptions, the two positions make comparable predictions. The delay-reduction hypothesis is also compared to several contemporary pscyhological accounts of choice. Results from several of our experiments with pigeons, designed as operant conditioning simulations of foraging, have shown the following: The more time subjects spend searching for or traveling between potential food sources, the less selective they become, that is, the more likely they are to accept the less preferred outcome; increasing time spent procuring (“handling”) food increases selectivity; how often the preferred outcome is available has a greater effect on choice then how often the less preferred outcome is available; subjects maximize reinforcement whether it is the rate, amount, or probability of reinforcement that is varied; there are no significant differences between subjects performing under different types of deprivation (open vs. closed economies). These results are all consistent with the delay-reduction hypothesis. Moreover, they suggest that the technology of the operant conditioning laboratory may have fruitful application in the study of foraging, and, in doing so, they underscore the importance of an interdisciplinary approach to behavior.

A representation is a remnant of previous experience that allows that experience to affect later behavior. This paper develops a metatheoretical view of representation and applies it to issues concerning representation in animals. To describe a representational system one must specify the following: the domain or range of situations in the represented world to which the system applies; the content or set of features encoded and preserved by the system; the code or transformational rules relating features of the representation to the corresponding features of the represented world; the medium, or the representation's physical instantiation; and the dynamics, or how the system changes with time. In part because of the behaviorist assumption that the hypothetical, covert changes occurring in an organism during learning correspond to the overt physical changes that are observed, issues of representation in animal behavior have been largely ignored as irrelevant or misleading. However, it can be inferred that representations, acting as models of environmental regularities, operate at many levels of behavioral functioning, both cognitive and noncognitive. Objections to the use of this concept in explanations of animal behavior, based on the claim that it is indeterminate and on behaviorist considerations of parsimony, can be answered. Animal representations may be specialized in terms of tasks and species. Data from tasks involving spatial memory, delayed matching-to-sample, and sequence learning suggest some foundations for a general theory of animal representations.

Previous lesion studies of color-reversal learning in pigeons show that an impairment results when (1) the tectofugal visual pathway is damaged at either the thalamic level (nucleus rotundus) or the telencephalic level (ectostriatum), or (2) the thalamofugal visual pathway is damaged at the telencephalic level (the visual Wulst). An impairment does not result, however, when the thalamic source of thalamofugal input (n. opticus principalis thalami or OPT) to the visual Wulst is damaged. These results suggest that the visual Wulst plays a role in color-reversal learning as a consequence of visual information routed from the tectofugal pathway via other visual areas in the telencephalon. One such area is the hyperstriatum ventrale (HV). In the present study, after ablation of the medial and lateral regions of HV, pigeons were trained postoperatively to discriminate between two colors presented simultaneously. After reaching criterion, the pigeons were required to perform a series of discrimination reversals in which the positive and negative stimuli were interchanged. Lesions of medial HV resulted in impaired performance of a color-discrimination task (i.e. original learning), but did not affect discrimination reversal. An impairment in color-reversal learning resulted from combined damage to lateral HV and the fronto-thalamic tract (FT), which carries ascending visual input from OPT to the visual Wulst. No deficits were observed when either lateral HV or FT were damaged alone. These findings suggest that both the thalamofugal and tectofugal pathways provide the visual Wulst with visual input relevant to color-reversal learning.

A serial-lesion technique was used to investigate interactions in visual processing between telencephalic components of the pigeon visual system. Pigeons were trained to discriminate pairs of stimuli that differed in color, intensity or pattern. After mastering the discrimination tasks, they were assigned to one of three groups. The first group (WI-EII) received lesions of the visual Wulst and were retested. After the discrimination tasks were again mastered, a second set of lesions was made, this time in the ectostriatum. The birds were tested once again after the second surgery. The second group (EI-WII), underwent the same sequence of events except that the order of the lesions was reversed. In the third group (E + W), lesions of both the visual Wulst and ectostriatum were made in a single operation, followed by retesting. The performance after the first lesion of the subjects in each of the two-stage lesion groups was typical of performance after such lesions; i.e. the birds with visual-Wulst lesions showed little or no impairment on any of the tasks, whereas the pigeons with ectostriatum lesions showed considerable deficits in intensity and pattern discrimination, which diminished after prolonged retraining. In contrast, the pigeons in the one-stage group (E + W) showed profound deficits that appeared to be permanent. The performance after the second operation of the WI-EII group was the same as that of pigeons with lesions of ectostriatum alone; i.e. destruction of ectostriatum first or second resulted in the same duration of impairment. The performance of the EI-WII group after its visual Wulst lesion, however, was similar to that observed in the E + W group. The results are interpreted as a reflection of parallel processing within the avian visual system; i.e. the presence of an intact tectofugal pathway may mask the effects of thalamofugal pathway interruption.

Nucleus rotundus and nucleus dorsolateralis posterior (DLP) are the thalamic components of two parallel pathways within the tectofugal division of the pigeon visual system. An earlier study (Hodos, Weiss & Bessette, 1986) had shown that lesions in direct telencephalic recipients of projections from rotundus and DLP produced postoperative elevations in size-difference thresholds only if the lesion included both structures. What was not revealed by their study was whether the integrity of both thalamic components is necessary for pigeons to discriminate small differences in the size of stimuli or whether the birds could still make the discrimination with only one of the two nuclei intact. This question was particularly important because no prior behavioral evidence existed to indicate that DLP plays a role in visual information processing. Therefore, 14 pigeons were tested preoperatively using a variant of the method of constant stimuli to determine the smallest difference between the size of two annuli that the subjects could discern. The comparison stimuli, which were presented in a successive discrimination procedure, ranged from 3.5–15 mm in diameter. After surgery, in which lesions were placed bilaterally in rotundus, DLP, or both structures, the subjects' size-difference thresholds were again determined. Combined lesions of rotundus and DLP resulted in impaired psychophysical performance. The postoperative behavior was characterized by initial elevations in threshold followed by a gradual improvement in performance. Some birds returned to their preoperative level. By comparison, subjects with lesions in rotundus or DLP alone showed an immediate return to their preoperative sensitivity level. These results indicate that both nuclei can process information about the size of visual stimuli. Moreover, the processing that occurs within either nucleus is sufficient for the pigeon to discriminate size differences. The present experiment provides the first behavioral evidence that DLP participates in visual information processing.

The performance of pigeons on a color-reversal learning task was assessed after thalamic lesions disrupting the thalamofugal and tectofugal visual pathways. Successful performance of a simultaneous color discrimination was accomplished after surgery, and a series of reversals of the original discrimination followed during which the positive and negative consequences associated with the stimuli were interchanged. Shimizu and Hodos (1989) had reported that lesions of two laminae in the visual wulst (IHA and HD), both targets of the avian thalamofugal pathway, resulted in increased errors in a color-reversal learning task in pigeons. This finding suggested that the thalamofugal pathway might play a role in visual discrimination involving stimulus context changes. In the present study, lesions of the OPT complex (the thalamic source of afferents to IHA and HD) were found to have no effect on color-reversal learning performance. Instead, we found that damage to nucleus rotundus (the thalamic component of the tectofugal pathway) resulted in deficits that were far in excess of those that had been obtained after IHA and HD lesions. We suggest that the color-reversal learning deficits after Wulst lesions are not due to the Wulst's connections with the thalamofugal pathway, but rather to its connections with the tectofugal pathway.

Tectal evoked potentials (TEPs) in response to sinusoidal gratings of different contrast, spatial and temporal frequency have been recorded from the tectal surface of the pigeon. Responses have been digitally filtered in order to isolate transient oscillatory (fast) potentials (50–150 Hz), transient slow potentials (1–50 Hz), and the steady-state second-harmonic component (16.6 Hz). Transient slow potentials, as well as the steady-state second-harmonic component, are band-pass spatially tuned with a maximum at 0.5 cycles/deg and attenuation at higher and lower spatial frequencies. The high spatial frequency cutoff is 4–5 cycles/deg. Both transient slow potentials and the steady-state second-harmonic component increase in amplitude as a function of log contrast and saturate at about 20% contrast. The contrast sensitivity, as determined by extrapolating TEP amplitude to 0 V is 0.1–0.2%. These characteristics of spatial-frequency selectivity and contrast sensitivity are similar to those reported for single tectal cells. Unlike slow potentials, oscillatory potentials are not band-pass spatially tuned. In addition, their contrast response function does not saturate at moderate contrast. These differences suggest that tectal evoked slow and fast potentials reflect the activity of different neuronal mechanisms.

Previous biochemical and immunohistochemical studies have shown that the neurotensin-related hexapeptide LANT6 is widespread and abundant in the avian nervous system. In the present study, immunohistochemical techniques were used to show that LANT6 is present in numerous cells of the retinal ganglion cell layer in pigeons. Consistent with the possibility that these LANT6+ retinal cells might be retinal ganglion cells, it was found that (1) the distribution of LANT6+ fibers and terminals in the central retinal target areas matched the distribution of central retinal projections; (2) the LANT6+ fibers and terminals are eliminated from retinal target areas by transection of the contralateral optic nerve; and (3) LANT6+ retinal cells in the ganglion cell layer can be retrogradely labeled by injections of fluorogold in the tectum. These results suggest that LANT6 may be utilized as a neuroactive substance by the central terminals of numerous retinal ganglion cells in birds. Similar anatomical findings have been previously reported for members of several other vertebrate groups, giving rise to the possibility that LANT6 (or its homologues in nonavians) may be a phylogenetically ubiquitous neuroactive substance used by retinal ganglion cells.

Columbidae (pigeons and doves) is one of the most threatened bird families in the world. I analysed data on the BirdLife International Species Information Database to examine the distribution and causes of threat among columbids. Of 304 species extant in the wild, 59 (19%) are threatened with extinction, 48 (83%) of which have restricted ranges. All but two threatened columbid species (97%) inhabit tropical forests, and of these, 45 are island species (78% of all threatened columbid species). The taxonomic distribution of columbids follows three coherent areas: the Americas; Europe, Africa, the Middle East and Central Asia; Asia, Australasia and Oceania. Asia, Australasia and Oceania support nearly two-thirds of all extant species and three-quarters of threatened species (44), most of which (84%) are restricted range insular species. Three countries within this area are the most important for the conservation of columbid diversity: Indonesia, the Philippines and French Polynesia. Together these three countries support 40% of extant species and half of all threatened species. The greatest causes of threat to columbids are (1) habitat loss and fragmentation due to agriculture and extraction, (2) hunting for food, and (3) alien predator species. Habitat loss and fragmentation are universal threats to columbids. Hunting, however, is a significantly greater threat to species in Asia, Australasia and Oceania than to species in the other two areas and urgently needs to be addressed. I discuss the conservation implications of these findings and make research recommendations to aid and encourage the conservation of threatened columbids and their habitats.

The avian ciliary ganglion (CG) controls choroidal blood flow by its choroidal neurons, and pupil constriction and accommodation by its ciliary neurons. It was previously reported that both choroidal and ciliary neurons label positively for NADPH diaphorase (NADPHd), a marker for nitric oxide synthase (NOS). To assess if this labeling is preganglionic or postganglionic and to determine if it is attributable to neuronal NOS (nNOS), we studied pigeon CG using NADPHd histochemistry and nNOS immunohistochemistry (IHC). Short-duration staining times by NADPHd histochemistry yielded intense labeling of structures that appeared to be the cap-like endings on ciliary neurons and the boutonal endings on choroidal neurons that arise from the nucleus of Edinger-Westphal (EW), and light or no postganglionic perikaryal staining. The light postganglionic staining that was observed tended to be localized to ciliary neurons. Consistent with this, NADPHd+ nerve fibers were observed in the postganglionic ciliary nerves but rarely in the postganglionic choroidal nerves. These same staining times yielded robust staining of neurons in the orbital pterygopalatine microganglia network, which are known to be nNOS+. Diffuse staining of CG perikarya was observed with longer staining durations, and this staining tended to mask the preganglionic labeling. Preganglionic NADPHd+ staining in CG with short staining times was blocked by the NOS inhibitors iodonium diphenyl (IDP) and dichlorophenol-indophenol (DPIP), but the diffuse postganglionic staining observed with the longer staining times was not completely blocked. Labeling of CG sections for substance P (SP) by IHC (which labels EW-originating preganglionic endings in CG) and subsequently for NADPHd confirmed that NADPHd was localized to preganglionic endings on CG neurons. Immunohistochemical double labeling for nNOS and SP or enkephalin further confirmed that nNOS is found in boutonal and cap-like endings in the CG. Two studies were then carried out to demonstrate that the nNOS+ preganglionic endings in CG arise from EW. First, NADPHd+ and nNOS+ neurons were observed in EW in pigeons treated with colchicine to enhance perikaryal labeling. Second, NADPHd+ and nNOS+ preganglionic endings were eliminated from CG ipsilateral to an EW lesion. These various results indicate that NOS is present in EW-arising preganglionic endings on choroidal and ciliary neurons in avian CG. NOS also appears to be found in some ciliary neurons, but its presence in choroidal neurons is currently uncertain.

Choroidal blood flow (ChBF) in birds is regulated by a neural circuit whose components are the retina, the suprachiasmatic nucleus, the medial division of the Edinger-Westphal nucleus (EWM), the ciliary ganglion, and the choriod. We have previously shown that lesions of EWM appear to result in pathological alterations in the retina. To determine whether EWM lesions also lead to altered visual functions, we have examined the effects of EWM lesions on visual acuity in pigeons. Bilateral lesions of EWM were made electrolytically, and visual acuity for high-contrast, square-wave gratings was determined behaviorally about 1 year later and compared to that of a group of pigeons that had received sham lesions of EW about 1 year prior to acuity testing. Because lesions targeting EWM invariably resulted in damage to the adjoining lateral part of the Edinger-Westphal nucleus (EWL), which controls pupillary constriction and accommodation, two additional control groups were studied. In one such control group, bilateral lesions in the area pretectalis (AP), which innervates the pupillary control part of EWL and thereby controls pupillary constriction, were made and the effects on visual acuity determined about 1 year later. In the second such control group, the effects of acute accommodative and pupillary dysfunction on acuity were studied in pigeons made cycloplegic. The accuracy of all lesions was later confirmed histologically. The mean acuities of birds with AP lesions (9.1 ± 1.4 cycles/deg) and sham lesions (7.1 ± 1.5 cycles/deg) were not significantly different from normal, based on published normative data on pigeons. In contrast, pigeons with lesions that completely destroyed EW bilaterally showed visual acuity (2.7 ± 0.1 cycles/deg) that was well below the acuity of the sham and AP-lesion control groups. The acuity of the cycloplegic pigeons (4.8 ± 0.3 cycles/deg) and one pigeon with a nearly complete bilateral EWL but a unilateral EWM lesion (6.4 cycles/deg) indicated that only about half of the loss with a bilateral EW lesion could be attributed to accommodative dysfunction. Thus, bilateral destruction of EWM appears to have led to a loss in visual acuity. This conclusion suggests that disruption of adaptive neural regulation of ChBF may impair visual function. Destruction of EWM was, however, associated with damage to the somatic components of the oculomotor and trochlear nuclei. The possibility cannot be excluded that such damage also contributed to the acuity loss.