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Color Learning in Honeybees
One of the most common ways that honey bees, Apis mellifera, demonstrate associative learning is in the context of color recognition and discrimination tasks. Just as vertebrate species such as mice or pigeons that can be trained to perform associative learning tasks, honey bees make excellent subjects for tasks involving discrimination and color memory. Beginning in the early 1900s, scientists Karl von Frisch and later Randolf Menzel began asking questions about the existence, learning rates, memory, and timing of color vision in bees.

Color Discrimination


The Austrian zoologist Karl von Frisch began the exploration of color vision in honey bees when he asked the first question in 1919: does color vision in bees exist? By making use of bees associative learning abilities he performed an elegant experiment to show that honey bees were in fact capable of color discrimination.

To test color vision, von Frisch first trained his honey bees to feed from a small dish filled with a nectar-like sugar water. This dish was placed over a small piece of blue colored cardboard so that the color was visible to the bees as they fed. Once the bees had become accustomed to the blue cardboard, von Frisch surrounded the blue piece of cardboard with other identically sized pieces in varying shades of gray and placed small dishes over each piece. If bees could not discriminate between colors, they would be unable to distinguish the blue piece from the many gray toned pieces. In the case that bees did not have color vision then, von Frisch predicted that the bees would visit the gray and blue pieces with equal frequency, as they would not be able to tell a difference between them.

When he allowed bees access to the dishes, however, he found that the vast majority of the bees flew directly to the blue piece of cardboard on which they had previously obtained their sugar-water reward. The bees largely ignored the gray pieces which had not been rewarded. This directed exploration and targeting of the blue cardboard showed the the bees could indeed discriminate between the gray and blue shades, showing that bees do possess color vision. Von Frisch repeated this same basic experiment to show that bees produced the same results with other colors like red and yellow. Later other researchers were able to apply this excellent experimental design to other vertebrates as well, making it an invaluable insight into testing color vision in many organisms.

Color Learning Rates and Preferences
After von Frisch’s initial studies, the German scientist Randolf Menzel continued the study of color vision in honey bees and performed more detailed tests. He was curious about which colors honey bees would be able to learn fastest and whether or not bees had a greater aptitude for learning certain colors.

He used lights of varying color and intensity to illuminate circles of light on a solid surface. This set up was similar to the pieces of colored cardboard employed by von Frisch, but by using light instead of cardboard, Menzel was able to change the intensity and color of light easily. He could simply adjust the projection of the light to create a wide variety of different experimental set-ups.



To test the intricacies of the bee color vision von Frisch had established, performed an experiment that aimed to test bees ability to distinguish between two different colors. To do this, Menzel used a projected circle of colored light surrounding a small dish that could hold a sugar-water reward. Menzel then projected a second circle of differently colored light surrounding a second dish some distance away from the first. Next, a single bee was placed equidistant between these two different lights and allowed to choose which dish to search for a sugar-water treat. Only one of the colored light circles surrounded a dish that contained sugar-water; the other was empty. Menzel was then able to measure how quickly the bees learned to preferentially search only the rewarded light and to ignore the dish surrounded by unrewarded light.

Interestingly, the results of the experiment showed that bees did not learn to discriminate between all color pairs equally well. The fastest rate of learning was when violet light was rewarded. The color that the bees had the most difficulty learning was green, and all other colors fell somewhere in between. This evidence of inherent bias is evolutionarily reasonable given that color vision in bees allows them to distinguish between different nectar-bearing flowers, much like the rewarded dishes. As more flowers are purple than green it makes sense that bees would be more sensitive to colors likely to result in nourishment.

Color Memory
After this work on color preferences, Menzel extended his experiments to study memory in honey bee color vision. He wanted to know how many trials were necessary for honey bees to reliably choose a previously rewarded color when presented with several choices for potential rewards and how long honey bees could retain information about which color would be rewarded.

To test these questions, Menzel performed a variety of experiments. First, he presented individual bees with a sugar reward on a colored background for just a single trial. He then kept these bees in small cages for several days without any further trials. After a few days, he presented each bee with several dishes each on a different colored background at once. One of the colors was the same as that used during the initial trial. The others were novel, unrewarded colors. Amazingly, after just one trial and several days without any exposure to the rewarded color, bees correctly chose to explore the color used in the first trial more than fifty-percent of the time.

Menzel then repeated this experiment with another group of bees, keeping all factors the same except that in the second round of testing he gave the bees three initial trials with the rewarded color instead of just one. After several days in confinement when the bees were presented with a choice of colors just as in the first experiment, they virtually always chose the color that had been used during the first three trials.

This ability to retain information about color-linked rewards over a period of several days and after only very minimal exposure to the colored background indicates the great strength of honeybees memory with respect to color vision.

Timing in Color Learning
The next big question was posed by one of Menzel’s students, Elizabeth Opfinger: when do bees register and learn color? They wanted to know if bees registered color before, during, or after receiving their sugar-water reward. In order to explore this intriguing question, Menzel and Opfinger changed the background color beneath a rewarded dish at different stages of the honey bee feeding process: the approach time, the feeding time, and the departure time.

The outcome of this experiment revealed that bees register color during both the approach and feeding stages of the exposure process. In order for a bee to accurately remember a given color, it must be present for approximately five seconds in total. Although it varies slightly, Menzel and his colleagues found that bees usually remember best when the stimulus is present for about three seconds during the approach and two seconds after landing and beginning to feed.

The Neurobiology of Color Vision


Color vision in honey bees can also be examined from a neurobiological perspective in terms of the structure and organization of their compound eyes.

In 1975 Menzel published a seminal paper describing the morphology and spectral sensitivity of the honey bee eye. He examined color coding the honey bee retina by using a technique to mark individual cells with a florescent dye and record from these cells as single units. Such fine structure analysis allowed him to determine that there are three types of receptors in the honey bee eye: 1) UV receptors, 2) blue receptors, and 3) green receptors. The three receptors are dominated by three rhodopsin-like pigments. These pigments have maximal absorbance at wavelengths corresponding to 350nm, 440nm, and 540nm.

As the cells were examined in detail, certain features were distinguishable for each type of receptor cell. UV cells were found to form the longest visual fibres. These long visual fibers penetrated the lamina with arborizations, a tree-like branching of the fibers and spines. Blue and green receptor cells have more shallow fibers.

Interestingly, Menzel found that most of the cells he studied had secondary sensitivities that corresponded to wavelength regions at which the other two receptor types were maximally active. He used spectral efficiency experiments to show that such corresponding wavelength receptivity is the result of electric coupling.