Odor Trail through Ant Country
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Odor Trail through Ant Country The desert ant’s use of its own built-in GPS – consisting of a sun-compass-based path integration system and visual landmarks – in locating its nest is a known phenomenon. Researchers recently ascertained, however, that this system also includes a sense of smell. Even more surprising is the discovery that these animals learn to distinguish between different odors in the nest environment, and use these like a map. Markus Knaden and his team at the Max Planck Institute for Chemical Ecology in Jena set out to search for clues in ant country.
FOCUS_Orientation A desert ant runs through the test channel to the feeder. The walls of the channel prevent the ant from seeing its environment. Under these test conditions, the ant can only navigate using its sense of smell. TEXT MARCUS ANHÄUSER K athrin Steck looks a bit tired. contained in the nest odor of desert they meet one another or arrive at the That’s not surprising – she has ants. Does every nest have its own nest, they touch one another extensive- been up all night. The biolo- odor? Or do all nests smell the same? ly with their antennae,” says Knaden. gist has been traveling since What odor emanates from the en- Once a foraging ant finds a food source, 2:00 a.m. Her trip started with trance, a roughly two-centimeter hole it informs its companions in the nest by a two-hour taxi ride through the dark- in the encrusted ground of a desiccated means of an attractant, a pheromone ness from the Tunisian town of Maha- salt lake? This is the typical habitat of trail on the ground. The ants follow this rès to Monastir. From there, she flew the subject of their research – the feisty, odor trail to the food source and then back to Germany, took a train from shiny black ant Cataglyphis fortis. carry the food back to the nest. The more Leipzig airport to Jena and, instead of animals use the trail, the more reinforced going home, went straight to the labo- A TRAIL OF ODORS FOR it becomes. The result is the familiar ant ratory at the Max Planck Institute for THEIR CONSPECIFICS trails, teeming with insects. Chemical Ecology. However, the Cataglyphis genus does The reason for the urgency is the For two weeks, Steck stood all day, every not deploy these types of ant trails at all. small glass of clear liquid in her back- day, in the blazing sunshine in temper- “They are lone warriors when it comes pack. It contains invisible odors ex- atures of up to 50 degrees Celsius. Bühl- to food,” says the behavioral ecologist. tracted from the nest entrance of a mann extracted the odors at the nest In the salt lake, only individual ants are desert ant. “These must be cooled as entrance while Steck pursued the long- seen flitting around, sometimes zigzag- Photo: MPI for Chemical Ecology much as possible and then quickly legged ants on their journeys across the ging around while foraging, sometimes processed,” says the 34-year-old post salty desert floor. hot-footing it toward the entrance hole doc, who works with Bill Hansson, an The olfactory orientation of ants has of the nest. “A pheromone trail would olfaction expert. long been established – even if it is not quickly disappear in the salt pan, where Kathrin Steck and her colleagues obvious to everyone. The ants pick up temperatures reach almost 60 degrees,” Cornelia Bühlmann and Markus Knaden the scent of their food and identify says Knaden. There are ant species that want to discover the substances that are friends and foes in ant country. “When do leave odor trails in such territories, 4 | 10 MaxPlanckResearch 17
FOCUS_Orientation » Cataglyphis occupies a very special place in the world of ant research. It is known as a “navigational genius.” “but these are mostly species that are ac- Over the decades, researchers discov- tive in the evenings or at night.” This ered that the insect actually uses sever- means that they engage in foraging al techniques to cross the salt pans in when temperatures cool down. search of rich pickings and return di- rectly to its nest. “They use a path in- DESERT ANTS WITH BUILT-IN GPS tegration system – based on a sun com- pass for measuring directions and a In addition, laying an odor trail is ben- step counter for measuring distances eficial only if there are plentiful food covered – and landmarks, to find the sources to exploit, that is, for ants who nest entrance,” says Knaden, summa- are carrying in those seeds that are ly- rizing the ants’ repertoire. Their facet ing on the ground behind a bush. eyes are thus bigger than those of However, Cataglyphis fortis ants have many other ants. They also have very a particular liking for dead insects. large, mobile antennae. And they carry these back to the nest Yet, until now, hardly anybody had in one piece. bargained on their sense of smell for Markus Knaden has spent 14 years guiding them across the hot, hostile researching Cataglyphis, a genus with salt flats. Not even Knaden. It was only approximately 60 species, of which 6 when he arrived in Bill Hansson’s de- can be found in the Sahara, its main partment in 2006 that the use of the habitat. Cataglyphis occupies a very Formicidae nose as a homing device special position in the world of ant re- came to his attention. “I had hoped methyl salicylate, decanal, nonanal search. It is known as a “navigational that the ants had learned not only vis- and indole. “We took environmental genius.” This is probably a slight exag- ual landmarks, but also possibly odor odors that are not connected with geration; many other animal species cues,” says the Max Planck scientist. food, but rather are produced, for ex- have comparable orientation skills, but Knaden and Steck first began system- ample, by plants and are highly vola- only a few of them are as well re- atically researching the substances that tile at high temperatures,” says Knaden. searched as the Cataglyphis. ants actually smell – or don’t smell. In a Food odors would have been too at- Since the 1960s, scientists have special laboratory such as the one oper- tractive. And the ants are not supposed been investigating how ants get their ated by Bill Hansson they have all the to be attracted by the odor – they are bearings in the desert. Back then, the equipment they need to systematically supposed to be able only to identify it father of Cataglyphis research, neuro- investigate odors. Steck tested a wide va- and remember it. physiologist Rüdiger Wehner from the riety of substances from the odor librar- University of Zurich, pitched his tents ies and, using ‘electroantennograms,’ RESEARCH ON DRY LAND in the Tunisian town of Maharès and demonstrated how the olfactory nerves began to examine the navigational abil- in the antennae respond to odors. The odor researchers then wanted to ity of these long-legged, elegant desert The two researchers then concen- find out whether the animals are capa- runners, which are roughly the same trated on four odors on which they ble of using these odors for orientation size as the common wood ants. trained the ants for their experiments: in the area around the nest entrance, 18 MaxPlanckResearch 4 | 10
above: Different strokes: The entrances to many of these nests consist of only an inconspicuous, thumbnail-size hole in the ground; other ants of the same species create striking nest mounds. One possible reason could be that returning ants use the mound as a landmark and therefore find their way back faster to the safety of the nest. right: Kathrin Steck marks individual ants. Using checklists, the researchers can see how often the individual animals have already Photos: MPI for Chemical Ecology (2) performed the relevant training. for example, in the same way that they use visual landmarks, such as a bush or a stone, to pinpoint the small entrance hole. The area studied is a desiccated salt lake in Tunisia, six kilometers long, three kilometers wide, about one hour 4 | 10 MaxPlanckResearch 19
above: The test site shows the salt pan in the background. Several such salt pans are dot- ted throughout Tunisia. This one measures approximately three by six kilometers. below: In temperatures of up to 50 degrees Celsius, Kathrin Steck stands in the blazing sunshine and tracks the ants’ search runs in the channel from the coast and the town of Maha- rès, where the scientists were staying. The mecca of Cataglyphis research. Rü- diger Wehner discovered the coastal area in 1968 and set up his base there. Markus Knaden, who completed his doctorate with Wehner and worked with him as a post doc, also began Photos: MPI for Chemical Ecology (2) working with desert ants in this area. When he moved to Bill Hansson’s department in Jena in 2006, he initial- ly had some bad luck: his ant popula- tion in Maharès was almost decimated. “We assume that it had something to do with the major use of pesticides in the nearby olive tree plantations,” says 20 MaxPlanckResearch 4 | 10
FOCUS_Orientation Large test fields painted on the ground allow the researchers to record the ants’ extensive search runs. This method was established by Rüdiger Wehner, the father of Cataglyphis research in the late 1960s, and remains a proven technique to this day. Knaden. He and Steck were thus forced ing ants. “We are the crazy people who el. The ants then have no clues as to search for a new population – and come every year with new equipment to where the hole is. Nevertheless, they found one at the said salt lake, 75 and a jeep,” says Knaden, smiling. they find it – because the nest has a kilometers away. Knaden saw the ad- The scientists set up their experi- particular smell. The researchers know vantage of their salt flat immediately ment in the salt lake, which contains this because each time the ants on the on their first visit: “The nests there were water only during the winter. For their panel reach the wind vane, which in- literally exploding with ants.” The first experiments, they needed only a dicates the wind direction in front nests were much bigger than those in round, five-millimeter-thick plastic and of the entrance, they turn and scurry Maharès, which is beneficial for carry- aluminum panel with a one-centimeter to the hole. On the leeward side, how- ing out spot checks and performing sta- hole in the middle, which was placed ever, they wander around, searching, tistical evaluations. over a nest entrance. Using this meth- even if they are only a few centimeters od, Bühlmann and Knaden were able to from their own home. A first impor- SWITCHING TO LANDMARK demonstrate that odor is actually a fac- tant finding. ORIENTATION tor in helping the ants find the en- trance when they return to the nest: FORAGING TRIP IN Even though research conditions are once the sun compass and path integra- THE ALUMINUM CHANNEL better at the salt pan, the scientists tor have brought the ant to within a from Jena don’t want to move there meter of the nest, the animal then For the subsequent, more complex Photo: MPI for Chemical Ecology lock, stock and barrel; their base camp switches to landmark orientation. “The tests, the field researchers needed alu- continues to be in Maharès. “The coast ant cannot actually see the hole itself, minum channels measuring two me- is simply nicer,” says Knaden. Better so it remembers a nearby stone or ters long and seven centimeters wide, climate, a fishing port, a few restau- bush,” says Knaden. U-shaped and open at the top, which rants. And the locals know the re- In the experiment, the researchers are placed on the ground. These chan- searchers, as they spend entire days can exclude such information by cov- nels are connected via a tube with an standing in the scorching heat, watch- ering the nest entrance with the pan- upturned bucket positioned over the 4 | 10 MaxPlanckResearch 21
top: The researchers paint several spots entrance hole to the Cataglyphis nest. ple, on the wall opposite the hole. of color on the abdomen of each ant The scientists use this test channel to They evidently use the spot as a land- so that they can distinguish between the ants on the training runs. hide all visual cues and the horizon mark and link this to the location of from the ants’ field of vision: “The an- the entrance. Would the same thing bottom: Markus Knaden and Kathrin Steck look at the experimental data on imals scuttle out of their nest under work with odors? the computer. the bucket. They go through the tube Because Knaden and Steck were not and arrive at the side wall of the alu- sure whether even one (and if so which) minum channel through a tiny hole. of the four odors identified in the lab- There they look for the food that we oratory – methyl salicylate, decanal, have laid out and come back to this nonanal and indole – could ultimately hole,” says Markus Knaden, describing be important, they drizzled the four the experiment. substances separately to form an odor Would the ants also be guided by square in the training channel in front an odor that does not emanate from of the entrance hole. The ants learned the hole itself, but originates from its very quickly that their entrance could environment? When Knaden and be found in this odor field. When they Steck examined this issue in their returned after their foraging trip in the channel experiment, they made an ex- aluminum channel, they just searched citing discovery that once again illus- inside the square for the entrance and trates the complex sensory abilities of found it immediately. these small insects. But something first had to go wrong before they realized A MAP OF VARIOUS ODORS the ants’ abilities. Photos: MPI for Chemical Ecology (2) The hole in the side wall of the alu- Unfortunately, the control experiments minum channel is so small that it is yielded no conclusive results: the re- very difficult for the insects to find un- searchers offered an odor field again, less it is marked. “It takes them forev- but this time without an entrance. The er to find it,” says Markus Knaden. ants did remember the odor and initial- They keep running by the hole. How- ly systematically frequented the odor ever, they find it immediately if the re- field, “but each time they hurried out searchers paint a black spot, for exam- again over the lines of the square,” says 22 MaxPlanckResearch 4 | 10
FOCUS_Orientation The Jena-based odor specialists set up camp in the Tunisian coastal town of Maharès. It has been the mecca for Cataglyphis research since 1968, when Swiss zoologist Rüdiger Wehner discovered the town and decided to use it as a base for his research. stink – is: carbon dioxide, or stale air. “The animals breathe in the nest, food is consumed there – all of this causes the level of CO2 in the air to rise,” ex- plains Knaden. That could account for the nest odor, but he isn’t sure. To find out exactly, Cornelia Bühl- mann and Kathrin Steck collected the nest odors mentioned at the start of this article using an aspirator. Here in Jena, Steck puts the odors back into the refrigerator, and now there is just one more thing to do: go home and go to bed. Knaden, describing the ants’ behavior. arate sensory inputs – as they do for see- And then the researcher realized they ing – which means that they rely on had made a serious error: “We had both of their antennae. According to GLOSSARY mixed up the odors. In the control Knaden, “Ants that only had one an- channel, they were in the wrong loca- tenna could no longer find their way Electroantennograms (EAG) tions in relation to where the entrance around.” This type of stereo smelling Electrophysiological discharge of so- hole was supposed to be.” among animals has been known for a called summating potential (super- imposed action potential of several The ants were visibly confused long time; researchers are sure that rats odor receptors) over the entire by this. When Kathrin Steck arranged and even humans use such a method. antenna when an odor is applied. the odors in the same order in the odor What they did not realize is that ants Path integrator square as they were in the test channel also have this facility. Desert ants use a path integration in front of the entrance hole, everything Yet a number of questions remain system. This consists of a compass, was back to normal again: the ants unanswered: Which landmarks are to determine where they are heading, and an odometer to measure the looked for the nest entrance only with- more important for the ants – visual distances they have traveled. The in the odor field as they had learned to or olfactory? And does the distribution compass uses a light phenomenon in do previously. “We had not expected the of odors at the nest entrance in the the sky that is invisible to the human odor array to be important for orienta- natural environment really play a role? eye and stretches across the entire celestial hemisphere as a large-scale tion purposes,” says Knaden. After all, the experiment with the polarization pattern. Cataglyphis The animals clearly navigate not aluminum channels is an artificial measures the distance it has traveled only on the basis of one or two odors in arrangement. It is possible that the by integrating the steps it has taken the mix and link these in their memory ants may use such an odor map less at on the journey. Rüdiger Wehner and with the entrance hole, but they actual- the nest entrance but more when for- his colleagues discovered this by arti- ficially lengthening or shortening the ly remember the individual locations of aging. A typical smell of home could length of the desert ants’ legs before the odor drops. Or, in a nutshell: Cat- also be enough for an ant to find its Photo: MPI for Chemical Ecology they returned to the nest from a feed- aglyphis smells in 3-D – a completely sur- own nest. er. The result was that animals manip- prising and conclusive result that nobody Following his latest field trip, ulated in this way over- or underesti- mated the return distance by exactly had reckoned with after 40 years of re- Markus Knaden is somewhat skeptical the amount by which the animals had search on Cataglyphis’ navigational skills. about this: “It looks like there’s one modified their step length due to the In order to be able to orientate general nest smell,” he says. Steck, change in the length of their legs. themselves spatially in such an olfacto- Bühlmann and he already suspect what ry landscape, the animals need two sep- the smell – or perhaps one should say 4 | 10 MaxPlanckResearch 23
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