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Concept: Karl von Frisch


In the late 1930s, Karl von Frisch reported that semiochemicals released upon injury, act as alarm substances (Schreckstoff) in fish. In Ostariophysi species, club cells in the epidermis are believed to contain cues related to alarm substance; however, the function of club cells, primarily as reservoirs of alarm substance has been debated. Here, I describe an alarm response in the Japanese rice fish Oryzias latipes (medaka), a member of the order Beloniformes. The response to alarm substance (Schreckreaction) in medaka is characterized by bouts of immobility and an increase in cortisol levels within minutes of exposure to conspecific skin extract. Histological analysis, however, suggests that club cells are either rare or absent in the medaka epidermis. In addition to describing an uncharacterized behavior in a vertebrate popular for genetic and developmental studies, these results support the hypothesis that the primary function of epidermal club cells may be unrelated to a role as alarm substance cells. The existence of similar behavioral responses in two evolutionarily distant but well established laboratory models, the zebrafish and the medaka, offers the possibility of comparative analyses of neural circuits encoding innate fear.

Concepts: Psychology, Genetics, Model organism, Oryzias latipes, Oryzias, Austrian nobility, Karl von Frisch, Schreckstoff


More than 100 years ago, Karl von Frisch showed that honeybee workers learn and discriminate colors. Since then, many studies confirmed the color learning capabilities of females from various hymenopteran species. Yet, little is known about visual learning and memory in males despite the fact that in most bee species males must take care of their own needs and must find rewarding flowers to obtain food. Here we used the proboscis extension response (PER) paradigm to study the color learning capacities of workers and drones of the bumblebee, Bombus terrestris. Light stimuli were paired with sucrose reward delivered to the insects' antennae and inducing a reflexive extension of the proboscis. We evaluated color learning (i.e. conditioned PER to color stimuli) in absolute and differential conditioning protocols and mid-term memory retention was measured two hours after conditioning. Different monochromatic light stimuli in combination with neutral density filters were used to ensure that the bumblebees could only use chromatic and not achromatic (e.g. brightness) information. Furthermore, we tested if bees were able to transfer the learned information from the PER conditioning to a novel discrimination task in a Y-maze. Both workers and drones were capable of learning and discriminating between monochromatic light stimuli and retrieved the learned stimulus after two hours. Drones performed as well as workers during conditioning and in the memory test, but failed in the transfer test in contrast to workers. Our data clearly show that bumblebees can learn to associate a color stimulus with a sugar reward in PER conditioning and that both workers and drones reach similar acquisition and mid-term retention performances. Additionally, we provide evidence that only workers transfer the learned information from a Pavlovian to an operant situation.

Concepts: Psychology, Honey bee, Bee, Bumblebee, Pollinators, Bombus terrestris, Bumblebees, Karl von Frisch


Female honeybees use the ‘waggle dance’ to communicate the location of nectar sources to their hive mates. Distance information is encoded in the duration of the waggle phase (von Frisch, 1967). During the waggle phase the dancer produces trains of vibration pulses, which are detected by the follower bees via Johnston’s organ located on the antennae. To uncover the neural mechanisms underlying the encoding of distance information in the waggle dance follower, we investigated morphology, physiology, and immunohistochemistry of interneurons arborizing in the primary auditory center of the honeybee (Apis mellifera). We identified major interneuron types, DL-Int-1, DL-Int-2, and Bilateral DL-dSEG-LP, that responded with different spiking patterns to vibration pulses applied to the antennae. Experimental and computational analyses suggest that inhibitory connection plays a role in encoding and processing the duration of vibration pulse trains in the primary auditory center of the honeybee.SIGNIFICANCE STATEMENTThe waggle dance represents a form of symbolic communication used by honeybees to convey the location of food sources via species-specific sound. The brain mechanisms used to decipher this symbolic information are unknown. We examined interneurons in the honeybee primary auditory center and identified different neuron types with specific properties. The results of our computational analyses suggest that inhibitory connection plays a role in encoding waggle dance signals. Our results are critical for understanding how the honeybee deciphers information from the sound produced by the waggle dance and provide new insights regarding how common neural mechanisms are used by different species to achieve communication.

Concepts: Brain, Species, Auditory system, European honey bee, Honey bee, Beekeeping, Bumblebee, Karl von Frisch


Just as biologists have their favored places for doing research, so do historians. As someone who likes working in archives, the most surprising thing the present author ever found was a particular letter that had been written to him by the ethologist Niko Tinbergen-but that Tinbergen had never sent. The letter included a detailed critique of the intellectual style and conceptual shortcomings of Tinbergen’s career-long friend and colleague Konrad Lorenz. The present author first saw the letter 3 years after Tinbergen’s death and 10 years after the letter was composed. Here we discuss the contents and historical context of that letter.

Concepts: Ethology, Richard Dawkins, Konrad Lorenz, Nikolaas Tinbergen, Karl von Frisch


In the 19(th) century, it was found that attraction of bees to light was controlled by light intensity irrespective of colour, and a few critical entomologists inferred that vision of bees foraging on flowers was unlike human colour vision. Therefore, quite justly, Professor Carl von Hess concluded in his book on the Comparative Physiology of Vision (1912) that bees do not distinguish colours in the way that humans enjoy. Immediately, Karl von Frisch, an assistant in the Zoology Department of the same University of Münich, set to work to show that indeed bees have colour vision like humans, thereby initiating a new research tradition, and setting off a decade of controversy that ended only at the death of Hess in 1923. Until 1939, several researchers continued the tradition of trying to untangle the mechanism of bee vision by repeatedly testing trained bees, but made little progress, partly because von Frisch and his legacy dominated the scene. The theory of trichromatic colour vision further developed after three types of receptors sensitive to green, blue, and ultraviolet (UV), were demonstrated in 1964 in the bee. Then, until the end of the century, all data was interpreted in terms of trichromatic colour space. Anomalies were nothing new, but eventually after 1996 they led to the discovery that bees have a previously unknown type of colour vision based on a monochromatic measure and distribution of blue and measures of modulation in green and blue receptor pathways. Meanwhile, in the 20(th) century, search for a suitable rationalization, and explorations of sterile culs-de-sac had filled the literature of bee colour vision, but were based on the wrong theory.

Concepts: Insect, Light, Color, Color vision, Color theory, Apiology, Austrian nobility, Karl von Frisch


A century ago, in his study of colour vision in the honeybee (Apis mellifera), Karl von Frisch showed that bees distinguish between a disc that is half yellow, half blue, and a mirror image of the same. Although his inference of colour vision in this example has been accepted, some discrepancies have prompted a new investigation of the detection of polarity in coloured patterns. In new experiments, bees restricted to their blue and green receptors by exclusion of ultraviolet could learn patterns of this type if they displayed a difference in green contrast between the two colours. Patterns with no green contrast required an additional vertical black line as a landmark. Tests of the trained bees revealed that they had learned two inputs; a measure and the retinotopic position of blue with large field tonic detectors, and the measure and position of a vertical edge or line with small-field phasic green detectors. The angle between these two was measured. This simple combination was detected wherever it occurred in many patterns, fitting the definition of an algorithm, which is defined as a method of processing data. As long as they excited blue receptors, colours could be any colour to human eyes, even white. The blue area cue could be separated from the green receptor modulation by as much as 50°. When some blue content was not available, the bees learned two measures of the modulation of the green receptors at widely separated vertical edges, and the angle between them. There was no evidence that the bees reconstructed the lay-out of the pattern or detected a tonic input to the green receptors.

Concepts: Eye, Color, Yellow, Primary color, Color vision, Color theory, Color wheel, Karl von Frisch


Oswald Schwarz, a urologist from Vienna, was a scholar of Anton Ritter von Frisch and Hans Rubritius. As a physician during World War I, he was confronted with numerous bullet wounds to the spinal cord. In 1919, he completed his professorial thesis"Bladder dysfunction as a result of bullet wounds to the spinal cord". Oswald Schwarz was known as a committed surgeon. As an urologist he also treated patients with sexual dysfunction. Besides his practical and scientific urology-related work, he was also interested in psychology and philosophy. He held lectures on both subjects earning himself the nickname, the Urosoph. In the 1920s, Oswald Schwarz belonged to the inner circle of Alfred Adler, the founder of Individual Psychology, and was editor of the first psychosomatic textbook published in German, “Psychological origin and psychotherapy of physical symptoms” (1925). In addition, Schwarz wrote numerous articles and several books on sexual medicine. He also made many valuable contributions to the development of medical anthropology. Altogether, his work includes over 130 publications. Faced with the rise of fascism and National Socialism in Europe, Oswald Schwarz, who was of Jewish origin, emigrated to England in 1934. There he died in 1949. Unfortunately his scientific work has largely been forgotten. The aim of the following article is to remind us of his important contributions to the field.

Concepts: Psychology, Medicine, Erectile dysfunction, Sigmund Freud, World War II, World War I, Alfred Adler, Karl von Frisch


Despite convincing data collected by microspectrophotometry and molecular biology, rendering sharks colourblind cone monochromats, the question of whether sharks can perceive colour had not been finally resolved in the absence of any behavioural experiments compensating for the confounding factor of brightness. The present study tested the ability of juvenile grey bamboo sharks to perceive colour in an experimental design based on a paradigm established by Karl von Frisch using colours in combination with grey distractor stimuli of equal brightness. Results showed that contrasts but no colours could be discriminated. Blue and yellow stimuli were not distinguished from a grey distractor stimulus of equal brightness but could be distinguished from distractor stimuli of varying brightness. In addition, different grey stimuli were distinguished significantly above chance level from one another. In conclusion, the behavioural results support the previously collected physiological data on bamboo sharks, which mutually show that the grey bamboo shark, like several marine mammals, is a cone monochromate and colourblind.

Concepts: Experimental design, Eye, Vision, Color, Primary color, Color blindness, Color vision, Karl von Frisch


Since the demonstration of color vision in honey bees 100 years ago by Karl von Frisch, appetitive conditioning to color targets has been used as the principal way to access behavioral aspects of bee color vision. Yet, analyses on how conditioning parameters affect color perception remained scarce. Conclusions on bee color vision have often been made without referring them to the experimental context in which they were obtained, and thus presented as absolute facts instead of realizing that subtle variations in conditioning procedures might yield different results. Here, we review evidence showing that color learning and discrimination in bees are not governed by immutable properties of their visual system, but depend on how the insects are trained and thus learn a task. The use of absolute or differential conditioning protocols, the presence of aversive reinforcement in differential conditioning and the degrees of freedom of motor components determine dramatic variations in color discrimination. We, thus, suggest top-down attentional modulation of color vision to explain the changes in color learning and discrimination reviewed here. We discuss the possible neural mechanisms of this modulation and conclude that color vision experiments require a careful consideration of how training parameters shape behavioral responses.

Concepts: Psychology, Insect, Visual perception, Honey bee, Color vision, Apiology, Hermann von Helmholtz, Karl von Frisch


Karl von Frisch’s studies of bees' color vision and chemical senses opened a window into the perceptual world of a species other than our own. A century of subsequent research on bees' visual and olfactory systems has developed along two productive but independent trajectories, leaving the questions of how and why bees use these two senses in concert largely unexplored. Given current interest in multimodal communication and recently discovered interplay between olfaction and vision in humans and Drosophila, understanding multisensory integration in bees is an opportunity to advance knowledge across fields. Using a classic ethological framework, we formulate proximate and ultimate perspectives on bees' use of multisensory stimuli. We discuss interactions between scent and color in the context of bee cognition and perception, focusing on mechanistic and functional approaches, and we highlight opportunities to further explore the development and evolution of multisensory integration. We argue that although the visual and olfactory worlds of bees are perhaps the best-studied of any non-human species, research focusing on the interactions between these two sensory modalities is vitally needed.

Concepts: Cognition, Visual perception, Perception, Sensory system, Sense, Olfaction, Odor, Karl von Frisch