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Book ; Online: What does the honeybee see? And how do we know?

Horridge, Adrian

2009

Abstract: ... that bee vision is adapted to the recognition of places, not things. In this volume, Adrian Horridge also ...

Abstract	This book is the only account of what the bee, as an example of an insect, actually detects with its eyes. Bees detect some visual features such as edges and colours, but there is no sign that they reconstruct patterns or put together features to form objects. Bees detect motion but have no perception of what it is that moves, and certainly they do not recognize "things" by their shapes. Yet they clearly see well enough to fly and find food with a minute brain. Bee vision is therefore relevant to the construction of simple artificial visual systems, for example for mobile robots. The surprising conclusion is that bee vision is adapted to the recognition of places, not things. In this volume, Adrian Horridge also sets out the curious and contentious history of how bee vision came to be understood, with an account of a century of neglect of old experimental results, errors of interpretation, sharp disagreements, and failures of the scientific method. The design of the experiments and the methods of making inferences from observations are also critically examined, with the conclusion that scientists are often hesitant, imperfect and misleading, ignore the work of others, and fail to consider alternative explanations. The erratic path to understanding makes interesting reading for anyone with an analytical mind who thinks about the methods of science or the engineering of seeing machines
Keywords	Science (General)
Size	1 electronic resource (359 p.)
Publisher	ANU Press
Document type	Book ; Online
Note	English ; Open Access
HBZ-ID	HT020087556
ISBN	9781921536991 ; 1921536993
Database	ZB MED Catalogue: Medicine, Health, Nutrition, Environment, Agriculture

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Article: Bee Vision is Totally Different

Horridge, Adrian

American bee journal

2018 Volume 158, Issue 1, Page(s) 65

Document type	Article
ZDB-ID	820372-6
ISSN	0002-7626
Database	Current Contents Nutrition, Environment, Agriculture

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Article: Parallel inputs to memory in bee colour vision.

Horridge, Adrian

Acta biologica Hungarica

2016 Volume 67, Issue 1, Page(s) 1–26

Abstract: 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, ... ...

Abstract	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.
MeSH term(s)	Animals ; Bees/physiology ; Color Perception ; Color Vision ; Memory ; Pattern Recognition, Visual
Language	English
Publishing date	2016-03
Publishing country	Hungary
Document type	Journal Article
ZDB-ID	803005-4
ISSN	1588-256X ; 0236-5383 ; 0001-5288
ISSN (online)	1588-256X
ISSN	0236-5383 ; 0001-5288
DOI	10.1556/018.67.2016.1.1
Database	MEDical Literature Analysis and Retrieval System OnLINE

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Article: How bees distinguish colors.

Horridge, Adrian

Eye and brain

2015 Volume 7, Page(s) 17–34

Abstract: Behind each facet of the compound eye, bees have photoreceptors for ultraviolet, green, and blue wavelengths that are excited by sunlight reflected from the surrounding panorama. In experiments that excluded ultraviolet, bees learned to distinguish ... ...

Abstract	Behind each facet of the compound eye, bees have photoreceptors for ultraviolet, green, and blue wavelengths that are excited by sunlight reflected from the surrounding panorama. In experiments that excluded ultraviolet, bees learned to distinguish between black, gray, white, and various colors. To distinguish two targets of differing color, bees detected, learned, and later recognized the strongest preferred inputs, irrespective of which target displayed them. First preference was the position and measure of blue reflected from white or colored areas. They also learned the positions and a measure of the green receptor modulation at vertical edges that displayed the strongest green contrast. Modulation is the receptor response to contrast and was summed over the length of a contrasting vertical edge. This also gave them a measure of angular width between outer vertical edges. Third preference was position and a measure of blue modulation. When they returned for more reward, bees recognized the familiar coincidence of these inputs at that place. They cared nothing for colors, layout of patterns, or direction of contrast, even at black/white edges. The mechanism is a new kind of color vision in which a large-field tonic blue input must coincide in time with small-field phasic modulations caused by scanning vertical edges displaying green or blue contrast. This is the kind of system to expect in medium-lowly vision, as found in insects; the next steps are fresh looks at old observations and quantitative models.
Language	English
Publishing date	2015-03-11
Publishing country	New Zealand
Document type	Journal Article
ZDB-ID	2587460-3
ISSN	1179-2744
ISSN	1179-2744
DOI	10.2147/EB.S77973
Database	MEDical Literature Analysis and Retrieval System OnLINE

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Article ; Online: How bees discriminate a pattern of two colours from its mirror image.

Horridge, Adrian

PloS one

2015 Volume 10, Issue 1, Page(s) e0116224

Abstract: 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 ... ...

Abstract	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.
MeSH term(s)	Animals ; Bees/physiology ; Color Perception ; Electrophysiology ; Pattern Recognition, Visual
Language	English
Publishing date	2015-01-24
Publishing country	United States
Document type	Journal Article
ZDB-ID	2267670-3
ISSN	1932-6203 ; 1932-6203
ISSN (online)	1932-6203
ISSN	1932-6203
DOI	10.1371/journal.pone.0116224
Database	MEDical Literature Analysis and Retrieval System OnLINE

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Article: How bees distinguish patterns by green and blue modulation.

Horridge, Adrian

Eye and brain

2015 Volume 7, Page(s) 83–107

Abstract: In the 1920s, Mathilde Hertz found that trained bees discriminated between shapes or patterns of similar size by something related to total length of contrasting contours. This input is now interpreted as modulation in green and blue receptor channels as ...

Abstract	In the 1920s, Mathilde Hertz found that trained bees discriminated between shapes or patterns of similar size by something related to total length of contrasting contours. This input is now interpreted as modulation in green and blue receptor channels as flying bees scan in the horizontal plane. Modulation is defined as total contrast irrespective of sign multiplied by length of edge displaying that contrast, projected to vertical, therefore, combining structure and contrast in a single input. Contrast is outside the eye; modulation is a phasic response in receptor pathways inside. In recent experiments, bees trained to distinguish color detected, located, and measured three independent inputs and the angles between them. They are the tonic response of the blue receptor pathway and modulation of small-field green or (less preferred) blue receptor pathways. Green and blue channels interacted intimately at a peripheral level. This study explores in more detail how various patterns are discriminated by these cues. The direction of contrast at a boundary was not detected. Instead, bees located and measured total modulation generated by horizontal scanning of contrasts, irrespective of pattern. They also located the positions of isolated vertical edges relative to other landmarks and distinguished the angular widths between vertical edges by green or blue modulation alone. The preferred inputs were the strongest green modulation signal and angular width between outside edges, irrespective of color. In the absence of green modulation, the remaining cue was a measure and location of blue modulation at edges. In the presence of green modulation, blue modulation was inhibited. Black/white patterns were distinguished by the same inputs in blue and green receptor channels. Left-right polarity and mirror images could be discriminated by retinotopic green modulation alone. Colors in areas bounded by strong green contrast were distinguished as more or less blue than the background. The blue content could also be summed over the whole target. There were no achromatic patterns for bees and no evidence that they detected black, white, or gray levels apart from the differences in blue content or modulation at edges. Most of these cues would be sensitive to background color but some were influenced by changes in illumination. The bees usually learned only to avoid the unrewarded target. Exactly the same preferences of the same inputs were used in the detection of single targets as in discrimination between two targets.
Language	English
Publishing date	2015-10-05
Publishing country	New Zealand
Document type	Journal Article
ZDB-ID	2587460-3
ISSN	1179-2744
ISSN	1179-2744
DOI	10.2147/EB.S89201
Database	MEDical Literature Analysis and Retrieval System OnLINE

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Article: How bees distinguish black from white.

Horridge, Adrian

Eye and brain

2014 Volume 6, Page(s) 9–17

Abstract: Bee eyes have photoreceptors for ultraviolet, green, and blue wavelengths that are excited by reflected white but not by black. With ultraviolet reflections excluded by the apparatus, bees can learn to distinguish between black, gray, and white, but ... ...

Abstract	Bee eyes have photoreceptors for ultraviolet, green, and blue wavelengths that are excited by reflected white but not by black. With ultraviolet reflections excluded by the apparatus, bees can learn to distinguish between black, gray, and white, but theories of color vision are clearly of no help in explaining how they succeed. Human vision sidesteps the issue by constructing black and white in the brain. Bees have quite different and accessible mechanisms. As revealed by extensive tests of trained bees, bees learned two strong signals displayed on either target. The first input was the position and a measure of the green receptor modulation at the vertical edges of a black area, which included a measure of the angular width between the edges of black. They also learned the average position and total amount of blue reflected from white areas. These two inputs were sufficient to help decide which of two targets held the reward of sugar solution, but the bees cared nothing for the black or white as colors, or the direction of contrast at black/white edges. These findings provide a small step toward understanding, modeling, and implementing in silicon the anti-intuitive visual system of the honeybee, in feeding behavior.
Language	English
Publishing date	2014-10-31
Publishing country	New Zealand
Document type	Journal Article
ZDB-ID	2587460-3
ISSN	1179-2744
ISSN	1179-2744
DOI	10.2147/EB.S70522
Database	MEDical Literature Analysis and Retrieval System OnLINE

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Article ; Online: How bees discriminate a pattern of two colours from its mirror image.

Adrian Horridge

PLoS ONE, Vol 10, Iss 1, p e

2015 Volume 0116224

Abstract	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.
Keywords	Medicine ; R ; Science ; Q
Language	English
Publishing date	2015-01-01T00:00:00Z
Publisher	Public Library of Science (PLoS)
Document type	Article ; Online
Database	BASE - Bielefeld Academic Search Engine (life sciences selection)

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Article: Parallel inputs to memory in bee colour vision

Horridge, Adrian

Acta biologica Hungarica. 2016 Mar., v. 67, no. 1

2016

Abstract: 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, ... ...

Abstract	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.
Keywords	Apoidea ; color ; color vision ; entomologists ; flowers ; foraging ; humans ; light intensity ; memory ; receptors ; researchers
Language	English
Dates of publication	2016-03
Size	p. 1-26.
Publishing place	Akadémiai Kiadó
Document type	Article
ZDB-ID	803005-4
ISSN	1588-256X ; 0001-5288 ; 0236-5383
ISSN (online)	1588-256X
ISSN	0001-5288 ; 0236-5383
DOI	10.1556/018.67.2016.1.1
Database	NAL-Catalogue (AGRICOLA)

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bis Jg. 2021: Bestellungen von Artikeln über das Online-Bestellformular
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Article ; Online: Generalization in visual recognition by the honeybee (Apis mellifera): a review and explanation.

Horridge, Adrian

Journal of insect physiology

2009 Volume 55, Issue 6, Page(s) 499–511

Abstract: During a century of studies on honeybee vision, generalization was the word for the acceptance of an unfamiliar pattern in the place of the training pattern, or the ability to learn a common factor in a group of related patterns. The ideas that bees ... ...

Abstract	During a century of studies on honeybee vision, generalization was the word for the acceptance of an unfamiliar pattern in the place of the training pattern, or the ability to learn a common factor in a group of related patterns. The ideas that bees generalize one pattern for another, detect similarity and differences, or form categories, were derived from the use of the same terms in the human cognitive sciences. Recent work now reveals a mechanistic explanation for bees. Small groups of ommatidia converge upon feature detectors that respond selectively to certain parameters that are in the pattern: modulation in the receptors, edge orientations, or to areas of black or colour. Within each local region of the eye the responses of each type of feature detector are summed to form a cue. The cues are therefore not in the pattern, but are local totals in the bee. Each cue has a quality, a quantity and a position on the eye, like a neuron response. This summation of edge detector responses destroys the local pattern based on edge orientation but preserves a coarse, sparse and simplified version of the panorama. In order of preference, the cues are: local receptor modulation, positions of well-separated black areas, a small black spot, colour and positions of the centres of each cue, radial edges, the averaged edge orientation and tangential edges. A pattern is always accepted by a trained bee that detects the expected cues in the expected places and no unexpected cues. The actual patterns are irrelevant. Therefore we have an explanation of generalization that is based on experimental testing of trained bees, not by analogy with other animals. Historically, generalization appeared when the training patterns were regularly interchanged to make the bees examine them. This strategy forced the bees to ignore parameters outside the training pattern, so that learning was restricted to one local eye region. This in turn limited the memory to one cue of each type, so that recognition was ambiguous because the cues were insufficient to distinguish all patterns. On the other hand, bees trained on very large targets, or by landing on the pattern, learned cues in several eye regions, and were able to recognize the coarse configural layout.
MeSH term(s)	Animals ; Bees/physiology ; Pattern Recognition, Visual ; Vision, Ocular
Language	English
Publishing date	2009-04-29
Publishing country	England
Document type	Journal Article ; Review
ISSN	1879-1611
ISSN (online)	1879-1611
DOI	10.1016/j.jinsphys.2009.03.006
Database	MEDical Literature Analysis and Retrieval System OnLINE

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