Colour Discrimination Learning Research Articles

Brain morphology correlates of learning and cognitive flexibility in a fish species (Poecilia reticulata).

Determining how variation in brain morphology affects cognitive abilities is important to understand inter-individual variation in cognition and, ultimately, cognitive evolution. Yet, despite many decades of research in this area, there is surprisingly little experimental data available from assays that quantify cognitive abilities and brain morphology in the same individuals. Here, we tested female guppies (Poecilia reticulata) in two tasks, colour discrimination and reversal learning, to evaluate their learning abilities and cognitive flexibility. We then estimated the size of five brain regions (telencephalon, optic tectum, hypothalamus, cerebellum and dorsal medulla), in addition to relative brain size. We found that optic tectum relative size, in relation to the rest of the brain, correlated positively with discrimination learning performance, while relative telencephalon size correlated positively with reversal learning performance. The other brain measures were not associated with performance in either task. By evaluating how fast learning occurs and how fast an animal adjusts its learning rules to changing conditions, we find support for that different brain regions have distinct functional correlations at the individual level. Importantly, telencephalon size emerges as an important neural correlate of higher executive functions such as cognitive flexibility. This is rare evidence supporting the theory that more neural tissue in key brain regions confers cognitive benefits.

The Neural Signature of Visual Learning Under Restrictive Virtual-Reality Conditions.

Honey bees are reputed for their remarkable visual learning and navigation capabilities. These capacities can be studied in virtual reality (VR) environments, which allow studying performances of tethered animals in stationary flight or walk under full control of the sensory environment. Here, we used a 2D VR setup in which a tethered bee walking stationary under restrictive closed-loop conditions learned to discriminate vertical rectangles differing in color and reinforcing outcome. Closed-loop conditions restricted stimulus control to lateral displacements. Consistently with prior VR analyses, bees learned to discriminate the trained stimuli. Ex vivo analyses on the brains of learners and non-learners showed that successful learning led to a downregulation of three immediate early genes in the main regions of the visual circuit, the optic lobes (OLs) and the calyces of the mushroom bodies (MBs). While Egr1 was downregulated in the OLs, Hr38 and kakusei were coincidently downregulated in the calyces of the MBs. Our work thus reveals that color discrimination learning induced a neural signature distributed along the sequential pathway of color processing that is consistent with an inhibitory trace. This trace may relate to the motor patterns required to solve the discrimination task, which are different from those underlying pathfinding in 3D VR scenarios allowing for navigation and exploratory learning and which lead to IEG upregulation.

Visual learning in a virtual reality environment upregulates immediate early gene expression in the mushroom bodies of honey bees

Free-flying bees learn efficiently to solve numerous visual tasks. Yet, the neural underpinnings of this capacity remain unexplored. We used a 3D virtual reality (VR) environment to study visual learning and determine if it leads to changes in immediate early gene (IEG) expression in specific areas of the bee brain. We focused on kakusei, Hr38 and Egr1, three IEGs that have been related to bee foraging and orientation, and compared their relative expression in the calyces of the mushroom bodies, the optic lobes and the rest of the brain after color discrimination learning. Bees learned to discriminate virtual stimuli displaying different colors and retained the information learned. Successful learners exhibited Egr1 upregulation only in the calyces of the mushroom bodies, thus uncovering a privileged involvement of these brain regions in associative color learning and the usefulness of Egr1 as a marker of neural activity induced by this phenomenon.

Motion cues from the background influence associative color learning of honey bees in a virtual-reality scenario

Honey bees exhibit remarkable visual learning capacities, which can be studied using virtual reality (VR) landscapes in laboratory conditions. Existing VR environments for bees are imperfect as they provide either open-loop conditions or 2D displays. Here we achieved a true 3D environment in which walking bees learned to discriminate a rewarded from a punished virtual stimulus based on color differences. We included ventral or frontal background cues, which were also subjected to 3D updating based on the bee movements. We thus studied if and how the presence of such motion cues affected visual discrimination in our VR landscape. Our results showed that the presence of frontal, and to a lesser extent, of ventral background motion cues impaired the bees’ performance. Whenever these cues were suppressed, color discrimination learning became possible. We analyzed the specific contribution of foreground and background cues and discussed the role of attentional interference and differences in stimulus salience in the VR environment to account for these results. Overall, we show how background and target cues may interact at the perceptual level and influence associative learning in bees. In addition, we identify issues that may affect decision-making in VR landscapes, which require specific control by experimenters.

The link between relative brain size and cognitive ageing in female guppies (Poecilia reticulata) artificially selected for variation in brain size.

Cognitive ageing is the general process when certain mental skills gradually deteriorate with age. Across species, there is a pattern of a slower brain structure degradation rate in large-brained species. Hence, having a larger brain might buffer the impact of cognitive ageing and positively affect survival at older age. However, few studies have investigated the link between relative brain size and cognitive ageing at the intraspecific level. In particular, experimental data on how brain size affects brain function also into higher age is largely missing. We used 288 female guppies (Poecilia reticulata), artificially selected for large and small relative brain size, to investigate variation in colour discrimination and behavioural flexibility, at 4-6, 12 and 24months of age. These ages are particularly interesting since they cover the life span from sexual maturation until maximal life length under natural conditions. We found no evidence for a slower cognitive ageing rate in large-brained females in neither initial colour discrimination nor reversal learning. Behavioural flexibility was predicted by large relative brain size in the youngest group, but the effect of brain size disappeared with increasing age. This result suggests that cognitive ageing rate is faster in large-brained female guppies, potentially due to the faster ageing and shorter lifespan in the large-brained selection lines. It also means that cognition levels align across different brain sizes with older age. We conclude that there are cognitive consequences of ageing that vary with relative brain size in advanced learning abilities, whereas fundamental aspects of learning can be maintained throughout the ecologically relevant life span.

Sex differences in color discrimination and serial reversal learning in mollies and guppies.

Behavioral flexibility provides an individual with the ability to adapt its behavior in response to environmental changes. Studies on mammals, birds, and teleosts indicate greater behavioral flexibility in females. Conversely, males appear to exhibit greater behavioral persistence. We, therefore, investigated sex differences in behavioral flexibility in 2 closely related molly species (Poecilia latipinna, P. mexicana) and their more distant relative, the guppy P. reticulata by comparing male and female individuals in a serial, visual reversal learning task. Fish were first trained in color discrimination, which was quickly learned by all females (guppies and mollies) and all molly males alike. Despite continued training over more than 72 sessions, male guppies did not learn the general test procedure and were, therefore, excluded from further testing. Once the reward contingency was reversed serially, molly males of both species performed considerably better by inhibiting their previous response and reached the learning criterion significantly faster than their respective conspecific females. Moreover, Atlantic molly males clearly outperformed all other individuals (males and females) and some of them even reached the level of 1-trial learning. Thus, the apparently universal pattern of higher female behavioral flexibility seems to be inverted in the 2 examined molly species, although the evolutionary account of this pattern remains highly speculative. These findings were complemented by the observed lower neophobia of female sailfin mollies compared with their male conspecifics. This sex difference was not observed in Atlantic mollies that were observed to be significantly less distressed in a novel situation than their consexuals. Hypothetically, sex differences in behavioral flexibility can possibly be explained in terms of the different roles that males and females play in mating competition, mate choice, and reproduction or, more generally, in complex social interactions. Each of these characteristics clearly differed between the closely related mollies and the more distantly related guppies.

Guppies learn faster to discriminate between red and yellow than between two shapes

AbstractAnimal species are expected to evolve specialised cognitive abilities to solve the tasks that are critical for their fitness. The literature contains several examples of specialised cognitive abilities, but few regard fish. The guppy, Poecilia reticulata, is a freshwater fish in which females choose their mates based on colouration, and orange‐coloured fruits are important diet enrichments for both sexes. For these reasons, we expect that this species has evolved enhanced learning abilities in colour discrimination compared to other types of discrimination. The comparison between studies in which guppies were tested for colour discrimination and studies in which guppies were tested for shape discrimination seems to support this hypothesis, but direct testing is still lacking. We experimentally compared the learning performance of guppies trained in a red–yellow colour discrimination learning task and that of guppies trained in a shape discrimination learning task using the same, automated conditioning procedure. Guppies trained in the colour discrimination showed greater learning performance, which provides support to the hypothesis that guppies possess enhanced colour discrimination abilities. Moreover, we found that male guppies performed better than females in both shape and colour discrimination learning.

Does Fine Color Discrimination Learning in Free-Flying Honeybees Change Mushroom-Body Calyx Neuroarchitecture?

Honeybees learn color information of rewarding flowers and recall these memories in future decisions. For fine color discrimination, bees require differential conditioning with a concurrent presentation of target and distractor stimuli to form a long-term memory. Here we investigated whether the long-term storage of color information shapes the neural network of microglomeruli in the mushroom body calyces and if this depends on the type of conditioning. Free-flying honeybees were individually trained to a pair of perceptually similar colors in either absolute conditioning towards one of the colors or in differential conditioning with both colors. Subsequently, bees of either conditioning groups were tested in non-rewarded discrimination tests with the two colors. Only bees trained with differential conditioning preferred the previously learned color, whereas bees of the absolute conditioning group, and a stimuli-naïve group, chose randomly among color stimuli. All bees were then kept individually for three days in the dark to allow for complete long-term memory formation. Whole-mount immunostaining was subsequently used to quantify variation of microglomeruli number and density in the mushroom-body lip and collar. We found no significant differences among groups in neuropil volumes and total microglomeruli numbers, but learning performance was negatively correlated with microglomeruli density in the absolute conditioning group. Based on these findings we aim to promote future research approaches combining behaviorally relevant color learning tests in honeybees under free-flight conditions with neuroimaging analysis; we also discuss possible limitations of this approach.

The town bird and the country bird: problem solving and immunocompetence vary with urbanization

Adaptation to urban habitats presumably requires changes in cognitive, behavioral, and physiological traits enabling individuals to exploit new resources. It is predicted that boldness, reduced neophobia, and enhanced problem-solving and learning skills might characterize urban birds compared with their rural conspecifics, while exposure to novel pathogens might require an enhanced immunity. To test these predictions, we assessed problem solving, color discrimination learning, boldness, neophobia, and immunocompetence in the bullfinch Loxigilla barbadensis, a highly opportunistic and innovative endemic bird in Barbados, wild-caught from a range of differently urbanized sites. Birds from urbanized areas were better at problem solving than their rural counterparts, but did not differ in color discrimination learning. They were also bolder but, surprisingly, more neophobic than rural birds. Urban birds also had an enhanced immunocompetence, measured with the phytohemagglutinin antigen. Our study sheds light on the trade-offs acting on animals exposed to changing environments, particularly in the context of urbanization.

Colour discrimination and associative learning in hatchling lizards incubated at ‘hot’ and ‘cold’ temperatures

The ability of an animal to acquire, process and learn from information in their environment is thought to be fundamental to fitness. We currently have a poor understanding of the learning ability of young animals within the first few months of their life, the types of learning they use and the extent of their learning ability. Furthermore, an animal’s developmental environment, such as nest incubation temperature, may profoundly influence motor and cognitive skills. We first tested the ability of hatchling three-lined skinks (Bassiana duperreyi) incubated at ‘hot’ and ‘cold’ temperatures to solve an instrumental (motor) task before assessing their ability to learn colour associations in a multi-stage instrumental task, with a choice reversal. While 53 (88.3 %) lizards successfully completed the training phase, 14 (46.7 %) of the ‘hot’ incubated and none of the ‘cold’ incubated lizards successfully completed the instrumental task. Thirteen of these lizards rapidly learnt to discriminate colours, and this culminated in eight individuals successfully completing a choice reversal. Hatchling B. duperreyi demonstrated surprisingly rapid learning, and these results highlight the potentially important role of cognition during development and ultimately, in fitness.

Long Term Effects of Aversive Reinforcement on Colour Discrimination Learning in Free-Flying Bumblebees

The results of behavioural experiments provide important information about the structure and information-processing abilities of the visual system. Nevertheless, if we want to infer from behavioural data how the visual system operates, it is important to know how different learning protocols affect performance and to devise protocols that minimise noise in the response of experimental subjects. The purpose of this work was to investigate how reinforcement schedule and individual variability affect the learning process in a colour discrimination task. Free-flying bumblebees were trained to discriminate between two perceptually similar colours. The target colour was associated with sucrose solution, and the distractor could be associated with water or quinine solution throughout the experiment, or with one substance during the first half of the experiment and the other during the second half. Both acquisition and final performance of the discrimination task (measured as proportion of correct choices) were determined by the choice of reinforcer during the first half of the experiment: regardless of whether bees were trained with water or quinine during the second half of the experiment, bees trained with quinine during the first half learned the task faster and performed better during the whole experiment. Our results confirm that the choice of stimuli used during training affects the rate at which colour discrimination tasks are acquired and show that early contact with a strongly aversive stimulus can be sufficient to maintain high levels of attention during several hours. On the other hand, bees which took more time to decide on which flower to alight were more likely to make correct choices than bees which made fast decisions. This result supports the existence of a trade-off between foraging speed and accuracy, and highlights the importance of measuring choice latencies during behavioural experiments focusing on cognitive abilities.

A field test of behavioural flexibility in Zenaida doves (Zenaida aurita)

Animals' ability to adjust their behaviour when environmental conditions change can increase their likelihood of survival. Although such behavioural flexibility is regularly observed in the field, it has proven difficult to systematically quantify and predict inter-individual differences in free-living animals. We presented 24 Zenaida doves (Zenaida aurita) on 12 territories with two learning tests in their natural habitat in Barbados. The dove pairs showed high site fidelity and territoriality, allowing us to test individuals repeatedly while accounting for the effects of territorial chases and pair bonds on our learning measures. We used a foraging apparatus that enabled Zenaida doves to access seed, yet excluded other species, and measured doves' performance on colour discrimination and reversal learning tests. We found that (1) doves on all 12 territories passed the two tests; (2) mates within a pair were consistently solvers or scroungers; (3) sex, body condition and territorial chases did not consistently affect learning rates; (4) tameness was a significant negative predictor of learning to feed from the foraging apparatus and (5) scrounging within pairs seemed to facilitate learning. Our study presents a method to quantify intraspecific differences in behavioural flexibility in the field and relate these to individuals' physical and social traits.

Fast learning but coarse discrimination of colours in restrained honeybees

Colours are quickly learnt by free-moving bees in operant conditioning settings. In the present study, we report a method using the classical conditioning of the proboscis extension response (PER) in restrained honeybees (Apis mellifera), which allows bees to learn colours after just a few training trials. We further analysed how visual learning and discrimination is influenced by the quality of a stimulus by systematically varying the chromatic and achromatic properties of the stimuli. Using differential conditioning, we found that faster colour discrimination learning was correlated with reduced colour similarity between stimuli. In experiments with both absolute and differential conditioning, restrained bees showed poor colour discrimination and broad generalisation. This result is in strong contrast to the well-demonstrated ability of bees to finely discriminate colours under free-flight conditions and raises further questions about the temporal and perceptual processes underlying the ability of bees to discriminate and learn colours in different behavioural contexts.

Cross-modal effects on learning: a seismic stimulus improves color discrimination learning in a jumping spider

The production of multimodal signals during animal displays is extremely common, and the function of such complex signaling has received much attention. Currently, the most frequently explored hypotheses regarding the evolution and function of complex signaling focus on the signal and/or signaler, or the signaling environment, while much less attention has been placed on the receivers. However, recent studies using vertebrates suggest that receiver psychology (e.g. learning and memory) may play a large role in the evolution of complex signaling. To date, the influence of multimodal cues on receiver learning and/or memory has not been studied in invertebrates. Here, we test the hypothesis that the presence of a seismic (vibratory) stimulus improves color discrimination learning in the jumping spider Habronattus dossenus. Using a heat-aversion learning experiment, we found evidence for a cross-modal effect on color learning. Over a series of training trials, individuals exposed to a seismic stimulus jumped onto the heated color less frequently and remained there for less time than did individuals not exposed to a seismic stimulus. In addition, in a final no-heat test trial, individuals from the seismic-present treatment were more likely to avoid the previously heated color than were individuals from the seismic-absent treatment. This is the first study to demonstrate a cross-modal influence on learning in an invertebrate.

Teleosts: Simple Organisms? Complex Behavior

ZebrafishVol. 3, No. 2 Guest EditorialTeleosts: Simple Organisms? Complex BehaviorLaure Bally-CuifLaure Bally-CuifSearch for more papers by this authorPublished Online:5 Jun 2006https://doi.org/10.1089/zeb.2006.3.127AboutSectionsPDF/EPUB ToolsPermissionsDownload CitationsTrack CitationsAdd to favorites Back To Publication ShareShare onFacebookTwitterLinked InRedditEmail FiguresReferencesRelatedDetailsCited byInhibitory Avoidance and Color Discrimination Learning in Zebrafish15 September 2010Zebrafish behavioural assays of translational relevance for the study of psychiatric diseaseReviews in the Neurosciences, Vol. 22, No. 1Development of the Dopamine Systems in Zebrafish Volume 3Issue 2Jun 2006 InformationCopyright 2006, Mary Ann Liebert, Inc.To cite this article:Laure Bally-Cuif.Teleosts: Simple Organisms? Complex Behavior.Zebrafish.Jun 2006.127-130.http://doi.org/10.1089/zeb.2006.3.127Published in Volume: 3 Issue 2: June 5, 2006PDF download

Sound improves visual discrimination learning in avian predators.

Aposematic insects use warning colours to deter predators, but many also produce odours or sounds when attacked by a predator. One possible role for these additional components is that they promote the association between the warning colour and the non-profitability it signals, thus reducing the chance of future attacks from visually hunting predators. This experiment explicitly tests this idea by looking at the effects of sound on a visual discrimination task. Young domestic chicks were trained to look for food rewards under coloured paper cones scattered in an experimental arena. In a subsequent visual discrimination task, they learned to discriminate between rewarded and non-rewarded hats on the basis of colour. Half the chicks performed this task in silence, whilst the other half had a tone played when they attacked non-rewarded hats. The presence of the tone improved the speed of colour discrimination learning. This demonstrates that there could be a selective advantage for aposematic coloured insects to emit sounds when attacked, since avian predators will learn to avoid their coloration more quickly. The role of psychological interactions between signal components in receivers is discussed in relation to the evolution of multimodal displays.

Effects of telencephalic ablation on short-term memory and attention in goldfish

Two hypotheses regarding the functions of the teleost telencephalon (short-term memory and nonspecific arousal hypotheses) were examined by using a Y-maze training paradigm. A delayed reinforcement method, which allowed the separation of choice process (a process in which choice responses are evoked) and reward process (a process in which choice responses are reinforced), showed that normal fish can acquire clear learned responses to choice stimuli under different stimulus conditions between the choice process and the reward process, while telencephalon-ablated fish showed greatly impaired learning performance. Neither normal nor telencephalon-ablated fish could acquire learned responses to choice stimuli under neutral stimulus conditions in the reward process with respect to choice stimuli in the choice process. These results suggest that the telencephalon facilitates extratelencephalic short-term memory function essential for memory retention of choice stimuli and evoked choice responses until reinforcement, and support the supplementary function of the telencephalon suggested previously [Ohnishi, K., Telencephalic function implicated in food-reinforced colour discrimination learning in the goldfish, Physiol. Behav., 46 (1989) 707–712 and Savage, G.E., Temporal factors in avoidance learning in normal and forebrainless goldfish ( Carassius auratus), Nature, 218 (1968) 1168–1169]. In addition, it was shown that cue information in the reward process is very important for the fixation of short-term memory of choice stimuli and choice responses. Furthermore, telencephalon-ablated fish also showed clear visual aspect selection, as did normal fish, when they were reinforced to a visual compound stimulus containing heterogeneous aspects (pattern and colour). This result shows that the telencephalon-related arousal or attentional function is not critical for aspect selection in goldfish. It seems that visual aspect selection in goldfish is performed without paying telencephalon-related `selective attention' to an aspect.

Telencephalic function implicated in food-reinforced color discrimination learning in the goldfish

The effects of telencephalic ablation on the learning of color discrimination were studied in the goldfish ( Carassius auratus) to determine the role of the telencephalon in the learning. The performances of normal and telencephalonless animals were compared in relation to their instrumental training rates. The animals were trained to discriminate between blue and green stimulating patches presented simultaneously. Normal and telencephalonless animals which underwent ten trials per day were unable to discriminate between the stimuli, but animals which underwent thirty trials showed learning. In animals which underwent twenty trials, there was a clear difference between the performances of normal and telencephalonless animals: the normal ones were able to learn, but the telencephalonless ones showed a significantly impaired learning ability. These results suggest that the telencephalon is not essential for the learning, but is supplementary in that it facilitates integration of neural events in extratelencephalic areas that are necessary for the instrumental process involved in color discrimination learning.

Behavioral asymmetries in visual learning of young chickens

Two-week-old chicks were trained in a simultaneous color discrimination learning task using food reinforcement. They had to discriminate between two discs of a different color whose right-left position was alternated at random. Results showed that chicks made more errors when the nonreinforced color was placed on the left with respect to the animal's body rather than when it was placed on the right. When chicks were trained on a right-left discrimination with two discs of identical color the asymmetry disappeared. Results are discussed in relation to current evidence for functional lateralization in the avian brain.

Effects of kainic acid and radiofrequency lesions of the pulvinar on visual discrimination in the monkey

Monkeys with thermocoagulation or kainic acid lesions of the pulvinar and unoperated control monkeys were tested in two tasks: pattern discrimination retention and color discrimination learning in which the stimuli were located at the response sites or were separated spatially from them (S-R separation). The monkeys with kainic acid pulvinar lesions were mildly impaired in retention of the pattern discrimination, but were unimpaired in the color discrimination tasks with or without the S-R separation. The monkeys with thermocoagulation lesions, like monkeys with superior colliculus lesions in a prior study, were severely impaired in performing one of the color discrimination tasks with S-R separation. These findings suggest that: (a) the inferior pulvinar, unlike the superior colliculus, does not contribute to the performance of discriminations involving S-R separation; and (b) corticotectal projections traversing the pulvinar and destroyed by the thermocoagulation lesions are crucial to the performance of discriminations involving S-R separation. The results of an earlier experiment also suggested that interruption of corticotectal fibers passing through the pulvinar impairs performance in another task sensitive to superior colliculus lesions — spatial localization of light flashes. Thus, corticotectal projections may be crucial for the contribution of the colliculus to performance in a variety of visual tasks.