Odor Trail through Ant Country

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Odor Trail through Ant Country
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.
Odor Trail through 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,

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Odor Trail through Ant Country
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,

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Odor Trail through Ant Country
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

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Odor Trail through Ant Country
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

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Odor Trail through Ant Country
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

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Odor Trail through Ant Country
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

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Odor Trail through Ant Country
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

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Odor Trail through Ant Country Odor Trail through Ant Country
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