Polarized Light in Animal Vision: Polarization Patterns in Nature

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Springer Science & Business Media, Jan 12, 2004 - Medical - 447 pages
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While the human eye can practically cope only with two aspects of light, brightness and colour, for many animals polarization is a further source of visual information.

This fascinating phenomenon of polarization sensitivity is comprehensively treated by Horvath and Varju. Starting with a short introduction into imaging polarimetry - an efficient technique for measuring light polarization - various polarization patterns occurring in nature are presented. Among them are the polarizational characteristics of water surfaces, mirages and the underwater light field as well as the celestial polarization patterns affected by the illumination conditions of sunrise, sunset, clear or cloudy skies, moonshine and total solar eclipses.

The major part of the book is dedicated to the question: How can animals perceive and use the natural and artificial polarization patterns? Following a detailed compendium of the physiological basis of polarization sensitivity, several case studies of animal behaviour determined or influenced by polarization are presented. It is shown how arial, terrestrial and aquatic animals use the celestial and underwater polarization for orientation, e. g. how polarized light serves honeybees or ants as a compass. Further, it is explained how man-made objects affecting the natural optical environment may disorientate animals. For instance, as in the case where oil or glass surfaces, asphalt roads, or plastic sheets used in agriculture can be more attractive for water-seeking polarotactic insects than the water surface, and where mayflies lay their eggs on dry asphalt roads or cars.

  

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Contents

Polarimetry From PointSource to Imaging Polarimeters
3
12 Elements of the Stokes and Mueller Formalism of Polarization
8
13 Polarimetry of Circularly Unpolarized Light by Means of Intensity Detectors
9
14 PointSource Scanning and Imaging Polarimetry
10
16 Colour Coding and Visualization of Polarization Patterns
11
18 Polarizational Cameras
12
Polarization Patterns in Nature
13
SpaceBorne Measurement of Earthlight Polarization
15
Oil Reservoirs and Plastic Sheets as Polarizing Insect Traps
215
212 The Waste Oil Reservoir in Budapest as a Disastrous Insect Trap for Half a Century
219
2121 Surface Characteristics of Waste Oil Reservoirs
220
2122 Insects Trapped by the Waste Oil
221
2123 Behaviour of Dragonflies Above Oil Surfaces
222
213 DualChoice Field Experiments Using Huge Plastic Sheets
223
214 The Possible LargeScale Hazard of Shiny Black Anthropogenic Products for Aquatic Insects
227
Why Do Mayflies Lay Eggs on Dry Asphalt Roads? WaterImitating Horizontally Polarized Light Reflected from Asphalt Attracts Ephemeroptera
229

Skylight Polarization
18
32 Celestial Polarization Measured by Video Polarimetry in the Tunisian Desert in the UV and Green Spectral Ranges
19
Principal Neutral Points of Atmospheric Polarization
23
41 Video Polarimetry of the Arago Neutral Point of Skylight Polarization
25
42 First Observation of the Fourth Principal Neutral Point
27
24Hour Change of the Polarization Pattern of the Summer Sky North of the Arctic Circle
32
Polarization Patterns of Cloudy Skies and Animal Orientation
36
62 Continuation of the ClearSky Angle of Polarization Pattern Underneath Clouds
37
63 Proportion of the Celestial Polarization Pattern Useful for Compass Orientation Exemplified with Crickets
38
GroundBased FullSky Imaging Polarimetric Cloud Detection
41
Polarization Pattern of the Moonlit Clear Night Sky at Full Moon Comparison of Moonlit and Sunlit Skies
47
Imaging Polarimetry of the Rainbow
51
Which Part of the Spectrum Is Optimal for Perception of Skylight Polarization?
53
102 Why Do Many Insects Perceive Skylight Polarization in the UV?
56
1022 Was the UV Component of Skylight Stronger in the Past?
57
1023 Relatively Large Proportion of UV Radiation in Skylight?
59
1024 Mistaking Skylight for GroundReflected Light?
60
1026 Were UV Receptors Originally Skylight Detectors and only later Incorporated into the Evector Detecting System?
61
1028 In the Spectral and Intensity Domain the Celestial Band of Maximum Polarization Is less Pronounced in the UV than in the Blue
62
10210 Perception of Skylight in the UV Maximizes the Extent of the Celestial Polarization Pattern Useful for Compass Orientation under Cloudy Skies
64
103 Resolution of the UVSkyPol Paradox
68
104 EVector Detection in the UV also Maximizes the Proportion of the Celestial Polarization Pattern Useful for Orientation under Canopies
69
105 Analogy Between Perception of Skylight Polarization and Polarotactic Water Detection Considering the Optimal Spectral Range
71
107 Why Do Crickets Perceive Skylight Polarization in the Blue?
72
108 Concluding Remark
73
Polarization of the Sky and the Solar Corona During Total Solar Eclipses
74
111 Structure of the Celestial Polarization Pattern and its Temporal Change During the Eclipse of 11 August 1999
75
112 Origin of the Evector Pattern During Totality
78
113 Neutral Points of Skylight Polarization Observed During Totality
80
114 Origin of the Zenith Neutral Point During Totality
83
116 Imaging Polarimetry of the Solar Corona
85
ReflectionPolarization Pattern of the Flat Water Surface Measured by 180 FieldofView Imaging Polarimetry
88
Polarization Pattern of a Fata Morgana Why Aquatic Insects Are not Attracted by Mirages?
92
Polarizational Characteristics of the Underwater World
95
Circulary Polarized Light in Nature
100
152 Circulary Polarized Light Reflected from the Exoskeleton of Certain Arthropods
101
153 Circulary Polarized Light Emitted by Firefly Larvae
102
Polarized Light in Animal Vision
105
From Polarization Sensitivity to Polarization Vision
107
162 Polarization Sensitivity Polarization Vision and Analysis of Polarization Patterns
108
163 Functional Similarities Between Polarization Vision and Colour Vision
111
164 How Can Skylight Polarization Be Used for Orientation?
112
165 Possible Functions of Polarization Sensitivity
115
166 How Might Polarization Sensitivity Have Evolved?
116
167 Polarization Sensitivity of Rhabdomeric Invertebrate Photoreceptors
117
1671 Hypothetical Polarizing Ability of the Dioptric Apparatus
118
1673 Origin of High Polarization Sensitivity
121
1674 Origin of Low Polarization Sensitivity
122
1675 Rhabdomeric Twist and Misalignment and Their Functional Significance
123
1676 Ontogenetic Development of Photoreceptor Twist Outside the Dorsal Rim Area of the Insect Eye
124
1677 Characteristics of the Anatomically and Physiologically Specialized PolarizationSensitive Dorsal Rim Area in Insect Eyes
125
1678 PolarizationSensitive Interneurons in Invertebrates
128
169 Polarization Sensitivity in Plants
130
Polarization Sensitivity in Terrestrial Insects
131
172 Flies
143
1723 Musca domestica Calliphora erythrocephala Calliphora stygia and Phoenicia sericata
144
1724 Drosophila melanogaster
146
173 Ants
147
174 Crickets
156
1743 Gryllus bimaculatus
157
1744 Gryllus campestris
160
Butterflies and Moths
165
1751 Papilio xuthus
166
1753 Polarized Light Reflected from Butterfly Wings as a Possible Mating Signal in Heliconius cydno chioneus
169
177 Cockroaches
172
178 Scarab Beetles
173
179 Response of NightFlying Insects to Linearly Polarized Light
176
Polarization Sensitivity in Insects Associated with Water
178
181 Velia caprai
180
184 Waterstrider Gerris lacustris
181
185 Backswimmer Notonecta glauca
183
186 Dragonflies Odonata
188
187 Dolichopodids
191
188 Mayflies Ephemeroptera
192
1810 Insects Living on Moist Substrata or Dung
195
1811 Mosquitoes
197
MultipleChoice Experiments on Dragonfly Polarotaxis
199
How Can Dragonflies Discern Bright and Dark Waters from a Distance? The Degree of Linear Polarization of Reflected Light as a Possible Cue for D...
206
221 Swarming Behaviour of Mayflies Above Asphalt Roads
231
222 MultipleChoice Experiments with Swarming Mayflies
232
223 ReflectionPolarizational Characteristics of the Swarming Sites of Mayflies
234
224 Mayflies Detect Water by Polarotaxis
236
225 Comparison of the Attractiveness of Asphalt Roads and Water Surfaces to Mayflies
239
ReflectionPolarizational Characteristics of CarBodies Why Are WaterSeeking Insects Attracted to the Bodywork of Cars?
241
Polarization Sensitivity in Spiders and Scorpions
243
242 Scorpions
246
Polarization Sensitivity in Crustaceans
247
251 Mangrove Crab Goniopsis cruentata
249
253 Copepod Cyclops vernalis
250
254 Larvae of the Crab Rhithropanopeus harrisi
251
255 Larvae of the Mud Crab Panopeus herbstii
252
256 Grapsid Crab Leptograpsus variegatus
253
258 Grass Shrimp Palaemonetes vulgaris
255
259 Crab Dotilla wichmanni
257
2510 Water Flea Daphnia
259
2511 Mantis Shrimps
263
Polarization Sensitivity in Cephalopods and Marine Snails
267
2612 Squids
269
2613 European Cuttlefish Sepia officinalis
272
262 Marine Snails
274
PolarizationSensitive Optomotor Reaction in Invertebrates
276
272 Honeybees
277
274 Rose Chafers
278
276 Optomotor Response to Over and Underwater Brightness and Polarization Patterns in the Backswimmer Notonecta glauca
287
Polarization Sensitivity in Fish
293
281 Fish in Which PolarizationSensitivity Was Proposed
294
2812 Tropical Halfbeaks Zenarchopterus dispar and Zenarchopterus buffoni
295
2813 Halfbeak Fish Dermogenys pusilus
296
2814 Goldfish Carassius auratus
297
2815 African Cichlid Pseudotropheus macrophthalmus
299
2816 Anchovies Engraulis mordax and Anchoa mitchilli
300
2817 Rainbow Trout Oncorhyncus mykiss
301
2818 Juvenile Salmonid Fish Oncorhynchus tnykiss Oncorhynchus clarki clarki Oncorhynchus nerka and Salvelinus fontinalis
306
282 Fish with Debated Polarization Sensitivity and Fish in Which Polarization Insensitivity Was Proposed
307
2822 Common White Sucker Catostomus commersoni
308
283 Possible Biophysical Basis of Fish Polarization Sensitivity
309
2832 Embryonic Fissures in Fish Eyes and Their Possible Role in the Detection of Polarization
311
2833 Paired Cones as a Possible Basis for Polarization Sensitivity in Fish
312
28332 Proposed Basis for Polarization Sensitivity in Rainbow Trout due to Internal Reflection from the Membranous Partitions of Double Cones
314
Polarization Sensitivity in Amphibians
317
291 Tiger Salamander Ambystoma tigrinum
318
292 RedSpotted Newt Notophthalmus viridescens
320
293 Larval Bullfrog Rana catesbeiana
321
294 Proposed Mechanisms of Detection of Polarization in Amphibians
322
Polarization Sensitivity in Reptiles
324
302 Desert Lizard Uma notata
325
303 Sleepy Lizard Tiliqua rugosa
326
Polarization Sensitivity in Birds
328
311 Crepuscularly and Nocturnally Migrating Birds
330
3112 Northern Waterthrush Seiurus noveboracensis and Kentucky Warbler Oporornis formosus
331
3113 YellowRumped Warbler Dendroica coronata
332
3114 Blackcap Sylvia atrkapilla
334
3115 Savannah Sparrow Passerculus sandwichensis
335
312 DayMigrating Birds
340
313 Birds Which Might Be Polarization Insensitive or not Use Skylight Polarization in Their Migratory Orientation
341
3131 Debated Polarization Sensitivity in the Homing Pigeon Columba livia
342
31311 The Position of the Sun Hidden by Clouds Could also be Determined on the Basis of the Colour Gradients of Skylight Under Partly Cloudy C...
348
3132 European Robin Erithacus rubecula
349
3133 Pied Flycatcher Ficedula hypoleuca
350
314 Proposed Mechanisms of Avian Polarization Sensitivity
351
3142 A Model of Polarization Detection in the Avian Retina with Oil Droplets
353
Human Polarization Sensitivity
355
322 Boehm Brushes
361
PolarizationInduced False Colours
362
332 Polarizational False Colours Perceived by Papilio Butterflies
364
3322 PolarizationInduced False Colours Perceived by a Weakly PolarizationSensitive Retina
369
3323 ReflectionPolarizational Characteristics of Plant Surfaces
374
3324 Do PolarizationInduced False Colours Influence the Weakly PolarizationSensitive Colour Vision of Papilio Butterflies Under Natural Conditions?
376
333 Polarizational False Colours Perceived by a Highly PolarizationSensitive Retina Rotating in Front of Flowers and Leaves
377
334 Camouflage Breaking via PolarizationInduced False Colours and Reflection Polarization
378
335 Is Colour Perception or Polarization Sensitivity the more Ancient?
379
A Common Methodological Error Intensity Patterns Induced by Selective Reflection of Linearly Polarized Light from Black Surfaces
381
References
385
Subject Index
417
Colour Illustrations
425
Copyright

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About the author (2004)

Gabor Horvath received a doctoral fellowship in 1989 in the Biophysics Group of the Central Research Institute for Physics of the Hungarian Academy of Sciences (Budapest), where he developed a mathematical description and computer modelling of retinal comet-like afterimages.

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