Christmas Special: Celestial Navigation

I already lampooned the idea of a virgin birth last year, and while folk stories abound at this time of the year because of the Solstice, we’ll stick with the Christian myth once again this year. Specifically, the part about the three kings who follow the star to find the stable of Bethlehem. This type of navigation was popular (and still is if you like to do things old-skool) and effective. In this post, I’m going to tell you about other animals who can also use celestial clues to navigate. Not very blasphemous (cf. Appendix 1), but it’s the best I could think of.

The first thing that must be said is that there’s a critical difference between how humans use the stars and sun, and how other animals use them. Humans make a map (of the constellations, etc.). There has classically been a debate about whether other animals do this, but the evidence mostly points to them using the stars/Sun/Moon as compasses. This requires a hefty amount of memorisation – memorisation so powerful, that a honeybee only needs to see a small patch of sky (e.g. between clouds) to find out what direction she’s flying in (Wehner, 1989), and the recognition is instantaneous (Sakura et al., 2012).

Putting birds in planetaria is a good experiment to see if they’re sensitive to star patterns. Emlen (1967) found that by producing artificial constellations, he could predict how his indigo buntings would change their orientation. It’s tempting to say that they got confused due to a different star map, but in fact, they orient themselves along the North-South axis of the constellations, not on the positions of the stars (Wiltschko et al., 1987) – it’s a compass, not a map; although it must be said that they do afterwards learn a map, as shown by Emlen (1972). Ants and honeybees can also memorise the patterns of constellations and stars (Collett & Collett, 2002).

I already wrote about ants. Briefly, desert ants use light polarised by the atmosphere as a compass (Wehner, 1994); this ability was first discovered in honeybees by von Frisch (1949), who also discovered that the waggle dance of bees also contains information about the sky, with the angle between the motion and the vertical being equal to the angle between the direction of the resource/new nest and the sun’s position (von Frisch, 1948) – this is also why the bee dance language is so dynamic, the sun’s position changes all the time and so the dance has to be modified every time.

The polarisation patterns are a function of the sun’s position in the sky; chromatic contrast also plays a big role, and both of these features are used to determine what direction the ant is facing and in what direction it’s supposed to go. Many other insects, especially those that have to migrate across long distances (Shashar et al., 2005), can do this as well, and all use the same apparatus. Generally, only UV, blue and green polarisation patterns are used (Wehner, 1991).

The compound eyes of these insects contain a special area called the dorsal rim area (Homberg & Paech, 2002; diagram above from same source), populated by photoreceptors with a high sensitivity to polarised and UV light. They also connect to specific areas of the lamina and medulla of the optic lobe. These areas are in turn populated by specific neurons, the polarisation-sensitive interneurons, whose firing rates are dependent on the polarisation of the light (Homberg et al., 2011) – this serves to clean the signal and remove noise. They’ve been found in representatives of the Odonata, Orthoptera, Blattodea, Hemiptera, Coleoptera, Neuroptera, Hymenoptera, Lepidoptera, and Diptera. The system has best been studied with locust and cricket brains (Pfeiffer et al., 2005), because they’re large enough to allow practical electrophysiological study. In crickets, there are three types of these neurons, attuned to firing at e-vectors oriented 10°, 60° and 130° to the head (Labhart et al., 2001). Kinoshita et al. (2007) suggest that these neurons also code for the chromatic pattern in the sky.

Orient yourself with the above diagram from Pfeiffer et al. (2005). The polarisation-sensitive interneurons in the locust are housed in a specific neuropil connected to the medulla, the signal going to the anterior optic tubercle (Homberg, 2004). From there, other neurons take the signal to the lateral accessory lobe (Homberg et al., 2003), where it is finally processed by the central complex (Heinze et al., 2009), where a “map” forms from the differences in the intensity of the activity from the incoming neurons (Heinze & Homberg, 2007). The interpreted signal is then sent back to the lateral accessory lobe and redirected to the thoracic motor centers and translated into movement (Träger & Homberg, 2011).

In ants, the ocelli can also provide information on polarisation patterns (Fent & Wehner, 1985). Whether this is also the case in other insects isn’t known, but in general, it’s worth noting that the transmission of the information from ocelli is faster than from compound eyes (Parson et al., 2006), so it shouldn’t be surprising if it turns out ocelli play a larger role than the specialised dorsal rim area.

Tiger salamanders can also use polarisation patterns (Adler, 1976). I don’t know the details of the mechanism, though. Hawryshyn (1992) suggested that fish can also use polarisation patterns, Munro & Wiltschko (1995) proposed the same for birds. However, no conclusive proof has been given so far. Interestingly, cephalopods are sensitive to polarisation patterns, but it isn’t known whether they use them for navigational purposes. Wolf spiders use celestial clues as well (Papi, 1955), getting the information from their anterior median eyes (Magni et al., 1964), but I’m not sure how widespread it is in other spider families.

The polarisation pattern produced by the moon is used by nocturnal animals (Ugolini et al., 1999), even if it is a million times dimmer than the sun’s on even the best nights (Gál et al., 2001). An example is dung beetles (Dacke et al., 2003). A big worry nowadays is that excessive light pollution is messing around with such navigational systems (Horváth et al., 2009), since human lights, when they get scattered through the atmosphere, get polarised (Kerola, 2006). Reflections on glass or water also get polarised (Horváth et al., 2009). An effect has already been demonstrated in the navigation of European moths (Nowinszky & Puskás, 2011), and more research is underway to determine how serious the problem potentially is.

The position of the Sun in the sky can be seen by all animals with eyes. And that source of information is used by all sorts of animals. We’ll stick with the arthropods. It was shown as early as Santschi (1911) that ants (he studied Lasius niger) can use the sun’s position as a compass. Cockchafers make sure to fly towards the sun after laying their eggs (Couturier & Robert, 1958). Herrnkind (1972) explains that crustaceans also use the sun for orientation – this also applies to those crabs that spend time on land, e.g. hermit crabs (Vannini & Chelazzi, 1981). Surprisingly, other underwater animals can use the sun, e.g. fish (Leggett, 1977) – I’m not sure how they deal with the effects of refraction; most likely they’re acclimatised to it from birth (just as terrastrial animals are acclimatised to the refraction through air). It’s common across all vertebrates as well – mammals use it (Lüters & Birukow, 1963), as do reptiles (Adler & Phillips, 1985), including birds (Kramer, 1957).

Using the sun as a compass is fine and all – except that sometimes, the sun isn’t visible due to clouds. So there has to be some kind of backup system. Birds use their circadian rhythm to keep them in the same rough direction – they sense that the sun moves ~15° every hour and make sure they keep that calibration (Schmidt-Koenig, 1960). This control has been investigated at the molecular level in monarch butterflies by Zhu et al. (2008).

An alternative, or complement (no generalisations can be made, it depends on the taxon), is to use the magnetic field. Salmon can do this, when they’re migrating under ice and so can’t see the sun (Quinn & Brannon, 1982).

The colour of the sky is also useful. Freshly-hatched turtles, for example, use the colour of the sky to find the horizon, which is assumed to be where the ocean is.

Theoretically, it is possible that some animals calculate their orientation from the direction and length of shadows, but this has still not been demonstrated (it would make an interesting study though, even if the results will likely be negative).

Just to round the post off, some trivia about human use of stars for navigation. Thales, the prototypical absent-minded philosopher/professor (or so he was portrayed by Plato) and all around brilliant guy (sage, engineer, statesman, geometry, astronomy, he worked in all of them), is credited by Callimachus with having discovered the use of the Little Bear as a navigation aid, and celestial navigation was considered one of the critical practical crafts to know (as we can see in the Socratic Dialogs). Pytheas of Massilia (late 4th century BC) made detailed calculations for the position of the North Star for use in navigation. Even so, no sailors dared to go beyond sight of the coast in the Mediterranean except on the best starlit nights, when the Pleiades and Sirius (mostly) were visible. Abbé de St. Pierre (17th century) named navigation as one of the most important sciences. Celestial navigation came to a high-point with the Portuguese sailors and explorers who started exploring the coast of Africa, using Polaris at night and the Sun during the day – this was a far cry from the pathetic travellings around the Mediterranean Sea, since many more latitudes had to be travelled through.

Anyway, I wish all of you a Merry Christmas/Hannukah/Solstice/Monkey/whatever it is you celebrate. You’ll get a Happy New Year’s wish at the New Year’s post.

Appendix 1:

Some of the hilarious hate-mail I got for the parthenogenesis post from overly-sensitive people who can’t take facts or a joke.

  • how dare you insult our Holy Mother!!! rot in hell heathen [I like how you prefer Mary to English grammar. I’m sure she’d find it cute, except that she was probably an illiterate. Also, no way in hell (pun maybe intended?) she knew modern English.]
  • I think you have a profoundly darkened view of the world. The Christmas story is a METAPHOR, not meant to be taken literally. Your post wasn’t an insult to us, although I’m sure that’s all you wanted. It was an insult t your intelligence for thinking people take the Virgin Birth as anything more than a allegory. Good day, sir. [O RLY? You may hold a high view of your fellow theists, but here are some poll results showing that they really do believe in the ridiculous crap and don’t take them as metaphors. Also, my post was an educational one. If you noticed, 90% of it was about the biology of sexual reproduction, or did the mention of the Virgin Mary automatically make you blind? Also, I’m an astigmatic, but my retina functions just fine.]
  • You are a disgrace how can you spew such vile nonsense on the internet. I will pray for you. [Could you just send me money instead? ← Note that punctuation mark.]
  • Typical liberal leftie asshole, using science to indoctrinate everyone else into his atheistic immoral worldview. Fuck you. [I am a liberal, I am a leftie, and I am an asshole. But I have never mentioned politics in any of my posts here, and certainly not in the parthenogenesis post, my dear projector bulb friend. Also, I wasn’t using science to indoctrinate, I was stating scientific facts that lead to an obvious conclusion. Also, did you really miss that 90% of the post had nothing to do with the Virgin? Finally, no word said on morality in this blog or in that post, and I haven’t revealed anything about my worldview either, except maybe indirectly that I like to live in the real world, not in a make-believe fantastical world that makes some people feel better about themselves at the expense of others.]


Adler K. 1976. Extraocular photoreception in amphibians. Photochemistry and Photobiology 23, 275-298.

Adler K & Phillips JB. 1985. Orientation in a desert lizard (Uma notata): time-compensated compass movement and polarotaxis. Journal of Comparative Physiology A 156, 547-552.

Collett TS & Collett M. 2002. Memory use in insect visual navigation. Nature Reviews Neuroscience 3, 542-552.

Couturier A & Robert P. 1958. Recherches sur les migrations du Hanneton commun (Melolontha melolontha L.). Annales Epiphyties 3, 257-329.

Dacke M, Nilsson D-E, Scholtz CH, Byrne M & Warrant EJ. 2003. Animal behaviour: Insect orientation to polarized moonlight. Nature 424, 33.

Emlen ST. 1967. Migratory Orientation in the Indigo Bunting, Passerina cyanea: Part I: Evidence for Use of Celestial Cues. The Auk 84, 309-342.

Emlen ST. 1972. The Ontogenetic Development of Orientation Capabilities. In: Galler SR, Schmidt-Koenig K, Jacobs GJ & Belleville RE. Animal Orientation and Navigation.

Fent K & Wehner R. 1985. Ocelli: A Celestial Compass in the Desert Ant Cataglyphis. Science 228, 192-194.

Gál J, Horváth G, Barta A & Wehner R. 2001. Polarization of the moonlit clear night sky measured by full-sky imaging polarimetry at full Moon: Comparison of the polarization of moonlit and sunlit skies. Journal of Geophysical Research 106, 22647-22653.

Hawryshyn CW. 1992. Polarisation vision in fish. American Scientist 80, 164-175.

Heinze S & Homberg U. 2007. Maplike Representation of Celestial E-Vector Orientations in the Brain of an Insect. Science 315, 995-997.

Heinze S, Gotthardt S & Homberg U. 2009. Transformation of Polarized Light Information in the Central Complex of the Locust. The Journal of Neuroscience 29, 11783-11793.

Herrnkind WF. 1972. Orientation in shore- living arthropods, especially the sand fiddler crab. In: Winn HE & Bolla B (eds.). Behavior of Marine Animals, Vol. 1.

Homberg U. 2004. In search of the sky compass in the insect brain. Naturwissenschaften 91, 199-208.

Homberg U & Paech A. 2002. Ultrastructure and orientation of ommatidia in the dorsal rim area of the locust compound eye. Arthropod Structure & Development 30, 271-280.

Homberg U, Hofer S, Pfeiffer K & Sebhardt S. 2003. Organization and neural connections of the anterior optic tubercle in the brain of the locust, Schistocerca gregaria. The Journal of Comparative Neurology 462, 415-430.

Homberg U, Heinze S, Pfeiffer K, Kinoshita M & el Jundi B. 2011. Central neural coding of sky polarization in insects. Phil. Trans. R. Soc. B 366, 680-687.

Horváth G, Kriska G, Malik P & Robertson B. 2009. Polarized light pollution: a new kind of ecological photopollution. Frontiers in Ecology and the Environment 7, 317-325.

Kerola DX. 2006. Modelling artificial night-sky brightness with a polarized multiple scattering radiative transfer computer code. Monthly Notices of the Royal Astronomical Society 365, 1295-1299.

Kinoshita M, Pfeiffer K & Homberg U. 2007. Spectral properties of identified polarized-light sensitive interneurons in the brain of the desert locust Schistocerca gregaria. The Journal of Experimental Biology 210, 1350-1361.

Kramer G. 1957. Experiments on bird orientation and their interpretation. Ibis 99, 196-227.

Labhart T, Petzold J & Helbling H. 2001. Spatial integration in polarization-sensitive interneurones of crickets: a survey of evidence, mechanisms and benefits. The Journal of Experimental Biology 204, 2423-2430.

Leggett WC. 1977. The ecology of fish migrations. Annual Review of Ecology and Systematics 8, 285-308.

Lüters W & Birukow G. 1963. Sonnenkompaßorientierung der Brandmaus (Apodemus agraiusPall.). Naturwissenschaften 50, 737-738.

Magni F, Papi F, Savely HE & Tongiorgi I. 1964. Research on the structure and physiology of the eyes of a lycosid spider. II. The role of different pairs of eyes in astronomical orientation. Archives Italiennes de Biologie 102, 123-136.

Munro U & Wiltschko R. 1995. The role of skylight polarization in the orientation of a day-migrating bird species. Journal of Comparative Physiology A 177, 357-362.

Nowinszky L & Puskás J. 2011. Light trapping of Helicoverpa armigera in India and Hungary in relation with the moon phases. Indian Journal of Agricultural Research 82, 154-157.

Papi F. 1955. Astronomische Orientierung bei der Wolfspinne Arctosa perita (Latr.). Zeitschrift für vergleichende Physiologie 37, 230-233.

Parsons MM, Krapp HG & Laughlin SB. 2006. A motion-sensitive neurone responds to signals from the two visual systems of the blowfly, the compound eyes and ocelli. The Journal of Experimental Biology 209, 4464-4474.

Pfeiffer K, Kinoshita M & Homberg U. 2005. Polarization-Sensitive and Light-Sensitive Neurons in Two Parallel Pathways Passing Through the Anterior Optic Tubercle in the Locust Brain. Journal of Neurophysiology 94, 3903-3915.

Quinn TP & Brannon EL. 1982. The use of celestial and magnetic cues by orienting sockeye salmon smolts. Journal of Comparative Physiology A 147, 547-552.

Sakura M, Okada R & Aonuma H. 2012. Evidence for instantaneous e-vector detection in the honeybee using an associative learning paradigm. Proc. R. Soc. B 279, 535-542.

Santschi F. 1911. Observations et remarques critiques sur le mecanisme de l’orientation chez les fourmis. Revue Suisse de Zoologie 19, 303-338.

Schmidt-Koenig K. 1960. Internal Clocks and Homing. Cold Spring Harbor Symposia on Quantitative Biology 25, 389-393.

Shashar N, Sabbah S & Aharoni N. 2005. Migrating locusts can detect polarized reflections to avoid flying over the sea. Biology Letters 1, 472-475.

Träger U & Homberg U. 2011. Polarization-Sensitive Descending Neurons in the Locust: Connecting the Brain to Thoracic Ganglia. The Journal of Neuroscience 31, 2238-2247.

Ugolini A, Melis C, Innocenti R, Tiribilli B & Castellini C. 1999. Moon and sun compasses in sandhoppers rely on two separate chronometric mechanisms. Proc. R. Soc. B 266, 749-752.

Vannini M & Chelazzi G. 1981. Orientation of Coenobita rugosus (Crustacea: Anomura): A field study on Aldabra. Marine Biology 64, 135-140.

Von Frisch K. 1948. Gelöste und ungelöste Rätsel der Bienensprache. Naturwissenschaften 35, 12-23.

Von Frisch K. 1949. Die Polarisation des Himmelslichtes als orientierender Faktor bei den Tänzen der Bienen. Cellular and Molecular Life Sciences 5, 142-148.

Wehner R. 1989. The Hymenopteran Skylight Compass: Matched Filtering and Parallel Coding. The Journal of Experimental Biology 146, 63-85.

Wehner R. 1991. Visuelle Navigation: Kleinsthirn-Strategien. Verhandlungen der deutschen zoologischen Gesellschaft 84, 89-104.

Wehner R. 1994. The polarization-vision project: championing organismic biology. In: Schildberger K & Elsner N (eds.). Neural Basis of Behavioural Adaptation.

Wiltschko W, Daum P, Fergenbauer-Kimmel A & Wiltschko R. 1987. The Development of the Star Compass in Garden Warblers, Sylvia borin. Ethology 74, 285-292.

Zhu H, Sauman I, Yuan Q, Casselman A, Emery-Le M, Emery P & Reppert SM. 2008. Cryptochromes Define a Novel Circadian Clock Mechanism in Monarch Butterflies That May Underlie Sun Compass Navigation. PLoS Biology 6, e4.

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