What does the Honeybee See? And how do we Know?

What does the Honeybee See? And how do we Know?: A critique of scientific reason

ADRIAN HORRIDGE
Copyright Date: 2009
Published by: ANU Press
https://www.jstor.org/stable/j.ctt24hf1n
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  • Book Info
    What does the Honeybee See? And how do we Know?
    Book Description:

    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.

    eISBN: 978-1-921536-99-1
    Subjects: Zoology, General Science

Table of Contents

  1. Front Matter
    (pp. i-iv)
  2. Table of Contents
    (pp. v-vi)
  3. ABOUT THE AUTHOR
    (pp. vii-viii)
  4. PREFACE
    (pp. ix-x)
  5. ACKNOWLEDGMENTS
    (pp. xi-xii)
  6. INTRODUCTION
    (pp. xiii-xviii)

    Small in size but high among the wonders of nature, insects delight and amaze us with their skill in flight and their obvious ability to see. A pursued dragonfly will turn and twist but will not escape the one chasing behind, yet it can stop in an instant or catch a mosquito in midair. Unerringly, a bee will take the direct path to the hive and a hoverfly will stand still in the air with invisible vibrating wings. These actions are all controlled by vision.

    We know, however, that when an engineer constructs a visual system it requires a huge...

  7. CHAPTER SUMMARY
    (pp. xix-xxii)
  8. GLOSSARY
    (pp. xxiii-xxiv)
  9. 01 EARLY WORK BY THE GIANTS
    (pp. 1-18)

    Palaeolithic humans—with excellent vision and endless opportunity—must have examined honeybees busy at their work and wondered what the insects saw and what they were doing, just as somebody else, watching the humans in turn, wondered what on earth they were doing studying bees. For a social animal, it was important to know who was watching what.

    The powerful obstacle to understanding what bees see—and they obviously see something—is that the human mind reads itself into the minds of others, even into bees. We call it anthropomorphism. We imagine that the bees are seeing things. We see...

  10. 02 THEORIES OF SCIENTIFIC PROGRESS: HELP OR HINDRANCE?
    (pp. 19-38)

    We expect to have a problem in understanding what honeybees see because they have a tiny brain combined with a very wide view of the world: multum in parvo. We must draw conclusions from the way they behave. We reasonably expect that they detect only relatively simple parts of the scene, but at first we are unable to imagine how they see anything in a moving panorama. To make progress, we have to devise ways of asking questions of the bees so that logical conclusions can be made from the way they react. This chapter is about making firm conclusions....

  11. 03 RESEARCH TECHNIQUES AND IDEAS, 1950 ON
    (pp. 39-62)

    Several powerful new techniques became available as a result of wartime research. First, the electronic equipment developed for sonar and radar—particularly the availability of low-noise high-impedance amplifiers, the oscilloscope, tape recorder and automated cameras—encouraged a burst of effort to record everything of interest, particularly in the visual system. This was undertaken by Roeder at Tufts; Burtt and Catton at Newcastle; Parry and Pringle in Cambridge, in the United Kingdom; McCann at Cal Tech; Bishop and Keehn at the University of Southern California; Kuiper at Gröningen; and Kuwabara, Naka and Eguchi at Fukuoka. It started with only a few...

  12. 04 PERCEPTION OF PATTERN, FROM 1950 ON
    (pp. 63-84)

    There were many earlier descriptions of how particular insects responded to lights or ran towards dark holes or contrasting edges, and several useful summaries of this kind of behaviour, categorised into taxes, kineses and tropisms (Fraenkel and Gunn 1940). The movements that insects make when stimulated by light, however, say almost nothing about the mechanisms of processing. Similarly, the outstanding work by Baerends (1941) and van Beusekom (1948) in Holland showed that wasps recognised their nest site by the memorised configuration of landmarks in different directions relative to each other, but this was about performance, not mechanism. In the second...

  13. 05 THE RETINA, SENSITIVITY AND RESOLUTION
    (pp. 85-116)

    The combination of anatomy, optics and electrophysiology of the honeybee eye provides a splendid illustration of science in action and the way to figure out the mechanisms of processing in vision. It is a mature topic, with a wide variety of approaches—notably, optics, pigment movements, transduction, signal detection, successive arrays of nerve cells in parallel pathways, compromises between receptor sensitivity and resolution, distinctions between line-labelled channels and the interesting compromise between crowding-in more processing versus simplicity for speed of action.

    The compound eye is composed of numerous simple eyes, called ommatidia, which are arranged side by side at a...

  14. 06 PROCESSING AND COLOUR VISION
    (pp. 117-146)

    As we look deeper in the optic lobes, we find progressively altered maps of the visual panorama. Behind the retina there are three successive neural regions, crowded with dendrites and synapses, called respectively the lamina, medulla and lobula (Figures 6.1–3). The neuron cell bodies, with no electrical activity, form a thick coat around them. The regions are separated by tracts of axons and are the result of the growth of the eye at the edges, so that groups of local circuits are reduplicated side by side in columns to form successive arrays of dendrites in layers. From the retina...

  15. 07 PILOTING: THE VISUAL CONTROL OF FLIGHT
    (pp. 147-176)

    What a delight it is to watch insects go about their daily life on a summer day. Most obviously, a large male butterfly flutters by on regular patrol around its territory and suddenly it recognises a female of its own species. A group of hoverflies hovers in separate stations in a shady place between trees; bees move from one flower to another of the same kind; a large fly weaves from side to side as it dashes past; and along the footpath a dragonfly hunts for mosquitoes. They all appear to see quite well and, after centuries of discussion about...

  16. 08 THE ROUTE TO THE GOAL, AND BACK AGAIN
    (pp. 177-206)

    Besides piloting, bees steer towards particular goals. They are, after all, heading somewhere. One of the lessons from human navigation is that several mechanisms are used and any cue can be useful. While circumventing obstacles, landmarks must be remembered, the general direction must be maintained and irrelevant things must be ignored. In early studies of insect navigation, with a single effect attributed to a single cause, the bees′ use of several mechanisms in parallel led to confusion. As usual, positive results were explained by the first idea that came to mind and unsound theory was only slowly recognised, so controversies...

  17. 09 FEATURE DETECTORS AND CUES
    (pp. 207-248)

    This chapter traces the effort from 1990 to 2008 to identify and characterise the parallel pathways of feature detectors and cues at the heart of the mechanism of visual processing in the bee. Bees have a few different kinds of feature detectors in large arrays that respond to parts of parameters in the pattern. The features in the parameters are edges or areas of black or colour; that is all. The analysis has been done with patterns subtending 30–45º at the eye, so the responses are limited to a small part of each eye. The responses of each kind...

  18. 10 RECOGNITION OF THE GOAL
    (pp. 249-262)

    We can now return to the topic that caused Forel, Lubbock and Plateau so much trouble in the late nineteenth century: how bees locate and then recognise their destination. The roadblock to progress at that time was that the bees had already arrived at their destination, so they were as confused by the changes in the flowers and rewards as the researchers were about what the bees expected to find. Research since then has shown that bees navigate in the right direction for the correct distance using a variety of flexible mechanisms. They care little about exact appearances along the...

  19. 11 DO BEES SEE SHAPES?
    (pp. 263-282)

    When the human eye looks at an object, it is almost impossible to avoid seeing its shape. We cannot imagine how we would not see the shape. So it might be difficult for readers to accept the conclusions so far reached—that bees detect cues in simple patterns that they do not see, and remember the directions of cues that enable them to identify places.

    It is usually assumed that bees probably remember vague, crude or fuzzy shapes, and that, because they have eyes, the onus of proof lies with those who would show that they do not. The opposite,...

  20. 12 GENERALISATION AND COGNITIVE ABILITIES IN BEE VISION
    (pp. 283-306)

    For a century, there have been claims of something in bee vision more subtle than the coincidences of feature detectors and cues. Anthropomorphism—that is, the tendency to put human capabilities into the brains of the bees—was not openly supported, but cognition trickled down from work on higher animals.

    Under the general heading of cognition in vision, the oldest belief was that the bees really saw and remembered the spatial layout of patterns. Also, it was thought that bees generalised patterns that looked similar to them. More recently, it was proposed that bees recognised patterns as a whole, that...

  21. AFTERTHOUGHTS
    (pp. 307-310)

    Let us start at the small end, with the nerve cells. Electrophysiology is an attractive technique for mechanistic analysis. When successful, the data flow from the electrodes and amplifiers. To explain even a little of the behaviour, however, electrophysiology requires a thorough knowledge of the neuron anatomy. One can go only so far with these methods and then find that the different kinds of data do not easily fit together. Neurons are relatively simple, but their combinations are devilishly difficult to unravel. The bee has small neurons, as yet obscure neuron anatomy in detail and behaviour with too many interacting...

  22. SUMMARY OF THE MODEL OF BEES′ VISUAL PROCESSING
    (pp. 311-318)

    For a bee, the parameters in the external panorama display two types of features—areas and edges—which are processed separately and not reassembled. Apart from motion detectors, the peripheral units of vision are feature detectors of two types: intensity detectors that respond to areas and modulation and orientation detectors that respond to passing edges. The feature detector responses are summed by type and position in each local region of the eye to form several types of cues, each with the position of its average. This summation destroys the local pattern. The cues are within the bee and must therefore...

  23. BIBLIOGRAPHY
    (pp. 319-360)