1. The Flowers Stay Put, but the Sun Does Not
When a honey bee finds good flowers and flies home, she can perform a waggle dance on the vertical comb. The angle of her dance on the comb corresponds to the direction of the sun, so the sun works as her reference point[2].
Now the trouble. The angle between the sun and the flowers is not fixed. The sun rises roughly in the east and sets roughly in the west, so an hour from now the same flowers sit at a different angle from the sun. A bee that simply repeated the angle she measured in flight would slowly point the wrong way. To give a lasting message, she has to know how far the sun has moved, even though it is out of sight.
Researchers call the skill that solves this a time-compensated sun compass: using the sun to find a direction while correcting for its changing position through the day[3]. Correcting for the sun's movement takes an internal sense of the time of day.
2. Using the Sun as a Compass Takes a Clock
Think of a sundial. A shadow pointing left means one thing at nine in the morning and something else at three in the afternoon. Without knowing the time, the shadow tells you almost nothing about north or south. The sun's direction can be misleading in the same way: in the morning it is in the east, but later it has moved across the sky.
So if bees really do correct for the sun, they should have some way of telling the time of day. Scientists did test whether bees can do that, and the first experiments came earlier than the sun-compass idea itself.
3. A Student Trained Bees to Show Up at the Feeder on Time (1929)
Ingeborg Beling studied under Karl von Frisch, the bee scientist who later shared a Nobel Prize for work on animal behavior[5]. In her 1929 doctoral thesis she did the first experimental test of what German researchers called bees' Zeitgedächtnis, or "time memory"[1].
She marked individual bees and set out sugar water on a table, but only during one window of the day, lasting between 1.5 and 4 hours. After a few days of this, she ran a test day with no food at all and counted every visit[1].
The bees came anyway. Visits increased around the usual feeding time. At other hours the bees rarely visited. In one trial with three daily feeding times, all 19 marked bees turned up on the test day. She could also retrain bees to a new time, and a single day of training was enough to set up the memory[1].
That the bees were not simply following the smell of food or hunger pangs was backed up by other checks. Bees found food in their stomachs at every hour of the day, and bees could be trained to several feeding times in one day[1]. Beling's own idea for why bees might have this ability was that many flowers open and give nectar only at certain times, so remembering the hour saves wasted trips[1].
4. The Bees' Clock Kept Running Without Sunlight, and Not Quite on a 24-Hour Beat
Was the clock inside the bees, or were they reading some daily signal from the outside world? Beling and her fellow student Oskar Wahl tried to rule out the outside world one piece at a time[1].
Beling used a windowless room painted black, with steady light and steady temperature and humidity, and the bees still learned a feeding time. She also tried bees that had been raised in an incubator and never lived in the wild, and they could be trained just as well. Wahl ran a test in a salt mine, a place where almost no cosmic radiation reaches, to look for any unknown outside rhythm. And when he compared 23 °C (73 °F) with 31 °C (88 °F), the timing did not change, which was a clue that the clock is not just a chemical process speeding up when it is warm[1].
Two details fit the idea of an inner clock that is close to, but not exactly, a day long. Beling fed bees every 19 hours for 16 days, and they never learned that rhythm; after the last feeding, they showed a small peak of searching 24 hours later, not 19. Wahl could not train bees to a 48-hour gap either, as they kept turning up every 24 hours[1]. And in Beling's constant-light room the bees arrived earlier than the trained time, which suggests a clock that runs slightly faster than 24 hours when it gets no cues[1].
The most dramatic test came in the 1950s. Max Renner trained bees in Paris, in a special room with constant light, temperature, and humidity, and flew them to an identical room in New York in about 22 hours. The bees kept to their own rhythm of roughly 24 hours instead of switching to New York time[1].
Putting this together, the bee's time sense is an internal clock. Light and darkness can reset it, but it keeps going when they are taken away. Researchers now call this kind of clock a circadian clock[1].
5. Bees That Had Only Seen the Evening Sun Still Guessed Where the Morning Sun Would Be (1994)
A clock is half of the answer. The bee also needs to know the sun's route: east in the morning, high at noon, west in the evening. Does she learn this by watching, or does she arrive knowing something about it?
In 1994, Fred Dyer and Jeff Dickinson designed an experiment to separate those two ideas. They raised bees in an incubator and let them fly outside and see the sun only in the late afternoon. Then, on a cloudy day, they let the bees fly for the first time in the morning and early afternoon, and watched the angle of their waggle dances. Clouds hid the real sun, so the bees had to guess where it was[2].
All morning the bees behaved as though the sun were sitting nearly still, about 180 degrees away from the average direction they had seen it in on earlier evenings. From about local noon onward, they used the evening direction[2]. In other words, they had never seen a morning sun, but they danced as if they expected the sun to be somewhere very different from where it had been in the evening.
The authors concluded that honey bees are "innately informed" about the general pattern of the sun's movement, enough to build a mental picture of a part of the sun's path they had never seen[2]. This is one experiment on one species under specific conditions. It does not say that bees know everything from birth, since they may still fine-tune details from experience.
6. What Scientists Still Do Not Know About the Bee's Sun Correction
Two open questions remain. The first is how exact the correction is. One idea is a smooth correction that follows the sun minute by minute. Another is a "step" rule: assume the sun is in the east all morning, then switch to the west at midday. A 2023 modeling paper notes that this step rule has been reported in honey bees. The paper's computer model, which looked at migrating animals rather than bees, found that a rough correction can still work well. Compensating for the sun's position with 80% accuracy cut the distance traveled by only about 2% in its simulation[3]. That is a calculation about migration, not a measurement of bees.
The second question is how, inside the bee's head, the body clock connects to the sun compass. A 2015 study of the honey bee's "sky compass" network found nerve cells that could be a place where clock signals feed in. This is a hint from the wiring, and the paper's title calls it a "possible" site[4]. The same level of caution applies: it shows that the cells sit near each other, not that they actually pass messages.
So the solid parts are three: bees keep time, the clock keeps running in constant conditions, and they act as if they know the sun's path. The open parts are the exact rule for correcting the angle and the way the clock connects to the compass.
7. Try It: Record How a Shadow Moves Over a Day
You can watch the thing bees have to keep track of. On a sunny day, pick a fence post, lamppost, or a stick pushed into the ground, somewhere safe and away from roads. Do not look at the sun.
In the morning, at midday, and in the late afternoon, put a pebble where the shadow's tip lands and write down the time. Then look at how the pebbles line up. The sun is on the opposite side of the post from the shadow, so the line of pebbles shows the sun's path as a shadow's path, and you can see how far it moved in a few hours. A bee has to carry that movement in her head.
If you want to dig deeper, the review of Beling's work (source [1]) is a good place to start, since it tells the story of her experiments and what we know now. It is written for scientists, so it is a stretch, but the figures and the historical account are easier to follow.
Sources
Reading status: [1] was read as full text through Europe PMC (the first part, up to Renner's Paris to New York experiment; the rest was not read). [2] was read as an abstract only. [3] was read as the abstract, introduction, and the start of the methods. [4] was read as the title and the first part of the abstract. [5] was read in full. The Beling review reports Renner's experiment and the salt-mine test as part of the evidence for an internal clock; I did not read Renner's or Wahl's original papers.
- Ingeborg Beling and the time memory in honeybees: almost one hundred years of research (review article, 2024, Europe PMC / PubMed Central PMC10995049). https://pmc.ncbi.nlm.nih.gov/articles/PMC10995049/ (Beling's 1929 experiments, Wahl's 1932 follow-up, constant-room and temperature tests, 19-hour and 48-hour cycles, Renner's transatlantic test.)
- Dyer and Dickinson, "Development of sun compensation by honeybees: how partially experienced bees estimate the sun's course," Proceedings of the National Academy of Sciences 91(10), 4471-4474 (1994). https://pmc.ncbi.nlm.nih.gov/articles/PMC43807 (Abstract only: incubator-reared bees, cloudy-day test, the 180-degree finding, and the authors' conclusion.)
- The efficiency of varying methods and degrees of time compensation for the solar azimuth (modeling study, 2023, PubMed Central PMC10663790). https://pmc.ncbi.nlm.nih.gov/articles/PMC10663790/ (Definition of a time-compensated sun compass, step compensation reported in honey bees, and the 80% and 2% model result. A model of migration, not of bee dances.)
- Zeller et al., "Transmedulla Neurons in the Sky Compass Network of the Honeybee (Apis mellifera) Are a Possible Site of Circadian Input," PLoS ONE (2015). https://pmc.ncbi.nlm.nih.gov/articles/PMC4667876 (Only the title and the beginning of the abstract were read, so the claim here is kept to "a possible site.")
- The Nobel Prize in Physiology or Medicine 1973, NobelPrize.org. https://www.nobelprize.org/prizes/medicine/1973/summary/ (Von Frisch shared the 1973 prize with Konrad Lorenz and Nikolaas Tinbergen.)
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