Proc. Nat. Acad. Sci.

USA
Vol. 71, No. 3, pp. 966-970, March 1974

Olfactory Basis of Homing Behavior in the Giant Garden Slug, Limax maximus
(digitate ganglion/locomotion/orientation/terrestrial pulmonate)

ALAN GELPERIN
Department of Biology, Princeton University, PrincetQn, New Jersey 08540

Communicated by V. G. Dethier, November 14, 1973

Time lapse photography of slugs living in
ABSTRACT
an experimental enclosure shows that these animals can
return to a homesite from over 90 cm by a direct route.
Slime trail following and vision are not involved in this
behavior. In the presence of a low velocity wind, homing
occurs upwind. Surgical disconnection of the presumptive
olfactory apparatus (digitate ganglion) from the central
nervous system eliminates homing. Neurophysiological
recordings from the receptor surface associated with the
digitate ganglion and the olfactory nerve demonstrate
the olfactory function of the digitate ganglion. The olfactory acuity and capacity for directed locomotion via olfactory cues are also relevant to studies of slug feeding behavior, ecology, and learning ability.

in the soil and forage over an area extending at least 4.5

meters from the home (14). Time lapse photography of grey
field slugs, Agriolimax reticulatus, locomoting on an enclosed
soil surface shows that the animals often return to the same
hole in the soil from which they emerged earlier in the night
(15). The present work documents homing behavior in Limax
maximus and presents initial physiological investigation of
its sensory basis.
MATERIALS AND METHODS

The behavioral experiments were done on slugs confined to a

1.5 by 1.7-meter area of moist filter paper bounded by a 2inch wide border of crystalline NaCl. An inverted clay flower
pot with four semicircular notches cut in its lip, was centrally
located and served as the animal's daytime resting site. The
filter paper was kept moist by inverted water reservoirs. Food
items such as carrot (Daucus carota), potato (Solanum tuberosum). dog food (Ken-L-Ration), or mushroom (Boletus edulis)
were supplied in a petri dish at one corner of the arena. Fluorescent room lights provided illumination and were automatically controlled to produce a cycle of 12 hr of light and 12 hr
of darkness. In some experiments, a plastic covering was used
to shield the arena from air currents.
A 16-mm camera modified for time lapse operation and
equipped with a wide-angle lens was mounted vertically 2.7
meters above the experimental arena. A xenon bulb flash unit
with a flash duration of 2 msec was triggered synchronously
with the camera shutter. The charging capacitor in the flash
unit was changed from 300 uf to 40 Mf to produce the least
intense flash which would give a distinct image oln plus-X
negative film with the lens wide open. A framing rate of 4 per
minute was used. Typically the camera was activated from
1700 hr to 0900 hr the following day. No behavioral response
could be detected to the light pulse emitted by the flash unit.
More than 1600 hr of activity were filmed and analyzed using
an analytical projector.
The 25 slugs used in these experiments were Limax maximus
and included both locally collected animals and individuals
reared from eggs in the laboratory. No differences in homing
ability between these two categories of animals were observed.
Anatomical studies oln the optic tentacles were done using
standard histological techniques to prepare 8-,Am serial sections of whole tentacles stained with Mallory Heidenhain's
triple stain (16). Neural pathways in the digitate (= tentacular) ganglion were stained using axonal iontophoresis (17) to
introduce Co++ ions into axon.s. The tissue was then treated
with ammonium sulfide, dehydrated with ethanol, and cleared

"Simple" animals often reveal their possession of sophisticated

behavioral machinery when experimental questions are asked
in the proper context. This is nowhere better documented than
in studies of orientation. The sun-compass orientation mechanism of bees and ants (1), the ability of noctuid moths to
steer their flight path away from bats using two sense cells
(2), and the apparent use of hydrodynamic cues by migrating
lobsters (3) are examples of complex neural mechanisms which
became apparent when physiological experiments were done
in an ethological context. The present experiments on homing
in Limax maximus were undertaken to probe the complexity
of behavior possible in a preparation amenable to cellular
neurophysiological analysis (4).
Homing behavior has been documented in a wide variety of
molluscan species. Aristotle described the homing behavior
of limpets and experiments to date are still searching for the
sensory basis of this behavior (5, 6). Octopus can return to its
nest after forays covering considerable distances (7). Small
colonies of the intertidal pulmonate Onchidium nest in rock
crevasses and after a period of feeding away from the nest,
all members of a particular colony simultaneously return directly to their nest (8). Capture and release experiments have
shown that the garden snail, Helix pomatia, can return to
sites favorable for overwintering with an angular error of less
than 300 over distances up to 40 meters (9). The sea hare
Aplysia is diurnally active and returns to a particular location
in its tank of seawater to sleep (10).
The literature contains scattered suggestions of homing behavior among slugs. The ability of Limax maximus to show
homing behavior has been referred to anecdotally by Taylor
(11), Pilsbry (12), and Frdmming (13). Field observations of
the California banana slug, Ariolimax columbianus, suggest
that animals establish a homesite by excavating a depression
Abbreviation: EOG, electro-olfactogram.

966

Proc. Nat. Acad. Sci. USA 71

(1974)

Olfactory Basis of Slug Homing

B'

Ce

967

FIG. 1. Track of single slug during one night.

in basic methyl benzoate (18). Ganglia so treated were studied

and photographed as whole mounts.
Electrical recordings from the receptor surface of an optic
tentacle were made with saline-agar filled electrodes of tip
diameter 50-100 /Am connected to a neutralized input capacitance dc amplifier. Polyethylene suction electrodes were attached to the olfactory nerve after severing its central connections. Both types of signals were recorded relative to a
ground electrode in the saline bath. The signals were displayed on a multichannel oscilloscope and either photographed directly or recorded using an FM tape recorder.
The saline used had the following composition in mM: Na
70, K 2.5, Ca 3.4, Mg 0.8, Cl 81, glucose 0.6, Tris 50.
RESULTS
The animals spend the daylight hours in the dark and humid
environment provided by the homesite. With a latency varying from several minutes to several hours after lights off, they
emerge and move about the arena at speeds ranging between
0.069 cm/sec to 0.26 cm/sec. These travels bring them in contact with the salt barrier, the food dish, other slugs, and ultimately, the homesite. Periods of locomotor activity are interslpersed with periods of sleep, sexual activity, or feeding. The
return to the homesite is often quite direct and over virgin

territory.
A representative tracing of the travels of a slug about the

experimental arena is shown in Fig. 1. Three of the trips are

short and in the immediate vicinity of the pot. These short
trips predominate during the first night in the apparatus
when the pot and filter paper are clean. Two of the trips are
more extensive and have terminal segments which clearly
suggest a directed locomotion back to the homesite. All of the
activity occurred during the dark period; the animal made its
final return home 2.2 hr before light onset. We do not know
when the animal actually decided to return home, but the
final segments of the two long trips demonstrate the animal's
ability to return home in a direct I)ath from the outer limits
of the arena over previously untraveled territory.

FIG. 2. Homeward paths of several different slugs. Arrow

provides constant compass direction reference.

Fig. 2 presents 10 return Ipaths selected to represent the

variation in directness of homing observed in this study. The
maximum distance from which homing occurred was 93 cm,
the outer limit of the arena. The animals used varied in body
length from 7.5 to 16.5 cm, and inter-optic tentacle distance
varied from 1.5 to 2.5 cm. Fig. 2 also illustrates that the same
slug can use different paths home on successive nights (Paths
C, F, J) and that two slugs living together can use different
paths home on the same night (Paths A, I).
The nonrandom nature of homing was tested mathematically in the following way. A set of linear path segments with
origins close to the periphery of the arena was selected; only
those paths which were linear because the animal moved
along the salt barrier were excluded. For the path whose origin was closest to the home, the flower pot subtended 150 of
the horizon. I assume, for purposes of this test, that the animal
selects his direction of travel from a 1800 sector. This assumption yields a probability of contacting the home by chance of
15/180 or 8.3%. The sample of 41 linear path segments contained 13 (32%) which were homing paths, a clearly nonrandom distribution.
Experiments were then directed to the question of the sensory cue providing direction to the homeward path. The animals are not following slime trails home, although they can
follow slime trails and do so routinely to locate sexual partners. The use of visual cues is possible but unlikely. The optical system and fine structure of the slug eye suggest poor
visual acuity (19) and light was available for only 2 msec
every 15 sec. The use of vision in homing was tested in two
ways. During the dark period, slugs were removed from the
home, placed at the periphery of the arena, and allowed to
return home in complete darkness, which they did. Two animals in which the optic nerves were successfully sectioned

968

Physiology: Gelperin

Proc. Nat. Acad. Sci. USA 71 (1971)

FIG. 3. Distribution of homeward path directions relative to

direction of prevailing wind. Arrow indicates wind direction.

bilaterally also retained the ability to return home by a direct

path if removed from the home and placed at the periphery
of the arena.
Two kinds of experiments suggest that olfaction may be the
key stimulus for homing in these exl)eriments. If a very gentle
wind is set up across the arena, the animals show a distinct
tendency to move against the wind. Fig. 3 shows the distribution of 42 homeward paths in relation to the wind direction
over the app)aratus. Five times as many returns occur from
the two downwind sectors as from the two upwind sectors
(42%o versus 8%7). Application of the x2 one-sample test to the
distribution shown in Fig. 3 indicates a significant deviation
from the expectation that returns are uniformly distributed
around the home (P < 0.001). A distribution of returns based
on the hypothesis that the two downwind sectors receive between 4 and 8 times as many returns as the two upwind sectors is not significantly different from the observed distribution (0.05 < P < 0.1).
The digitate ganglion, located at the distal end of the
olfactory nerve in the dorsal or optic tentacles, has been
assigned an olfactory function based on behavioral (20-22)
and histological (23-25) observations. To test the idea that
the digitate ganglion is important in homing behavior, slugs
were subjected to bilateral olfactory nerve section. They were
anesthetized with CO2 and an incision made in the lateral
body wall of the head. The olfactory nerves were cut where
they emerge from the cerebral ganglia. Control slugs received
the same operative procedure except the nerves were not cut.
Operations were verified by autopsy. The six slugs successfully operated on, never homed again whereas the controls
continued to do so. The excursions of the operated animals
were of normal extent and at normal speed. However, even
though they occasionally passed within 2 cm of home, the
operated slugs did not return to it.
Several behavior patterns exhibited by normal slugs are
suggestive of an olfactory sensitivity. Locomotion of any tyele
is always accompanied by movements of the optic tentacles
which sweep through an arc of 15-20 on either side of the
midline. During homeward locomotion, animals often exhibit
a characteristic "head-waving" behavior during which the
anterior end of the body is lifted above the substrate and the
head moved from side to side with a frequency of approximately 1-2 per sec. Paths toward odorous foods, particularly
fungi, are also often quite direct over distances up to 80 cm.
The digitate ganglion is situated within the distal end of
the cylindrical tentacle retractor muscle. The finger-like
processes emanating from the ganglion innervate a distinctive

:.:K-

FIG. 4. Digitate ganglion after CoCl2 was iontophoresed

toward the ganglion via axons in the olfactory nerve and after
cobalt was precipitated with ammonium sulfide. Dark areas out
of focus at top are pigment cells in sensory epithelium. Three
neuronal somata in the ganglion are indicated by short arrows.
The majority of the fibers in the olfactory nerve terminate in the
ganglion. scale bar = 500 um.

el)ithelial lpad or sensory zone (26) at the end of the tentacle.

Presumptive sensory neurons located in and under the sensory

el)ithelium send lprocesses into the digitate ganglion, as do

large numbers of 5-7 Azm cells located in the distal extensions

of the ganglion (24). A mantle of neurosecretory cells surrounds the ganglion (27). Cobalt backfills of the olfactory
nerve reveal six to eight large cells in the body of the ganglion
and several fiber tracts ending within it (Fig. 4). This suggests
that in fact the majority of the axons in the olfactory nerve
are second order processes.
To test the olfactory function of the digitate ganglion
physiologically, a preparation of the sensory el)ithelium,
digitate ganglion, and olfactory nerve was isolated from the
animal and set up in vitro so that the sensory surface was exl)osed to the air while the ganglion and nerve were immersed
in saline. An agar-filled electrode recorded from the sensory
surface (28, 29) and a suction electrode monitored the olfactory nerve. Filter paper discs were saturated with odorants
and l)laced in a Swinny adapter ('Millipore Corp.) mounted
on a 1-cc syringe. The syringe was mounted so that the tip of
the adapter was I cm from the receptor surface. Stimuli were
delivered by hand.
A puff of air containing an odorant such as amyl acetate
elicits a large compound action l)otential in the olfactory
nerve (Fig. 5. A) and a negative electro-olfactogram (EOG)
(Fig. 5, A1,). Several small single unit responses are evident in

Proc. Nat. Acad. Sci. USA 71

(1974)

Olfactory Basis of Slug Homing

969

Al preceding and following the compound action potential.

The single unit responses typically lasted for the duration of
the active phase of the EOG. A puff of moist air produces no
response (Fig. 5, B1, B2). The EOG is not recorded from epithelium outside the sensory patch. The size of the compound
action potential and EOG can be increased by more rapid application of a given volume of odorant-laden air. Similar responses were obtained to aqueous extracts of carrot and potato. These data demonstrate the olfactory function of the
digitate ganglion.
DISCUSSION
The results presented here demonstrate the homing ability of
Limax maximus. The maximum distance over which Limax
can forage and still return home is unknown. Estimates of
this distance require knowledge of the chemical species used
as an olfactory cue and information on the behavioral threshold to this chemical (30). The behavioral threshold could be
very low since insect and vertebrate olfactory systems can
trigger a behavioral response to a few molecules of odorant
(30). If the initial behavioral response triggered by the odor
is movement ul)willd, a detectable gradient is unnecessary and
the odor will be effective in promoting homing even at the
fringes of its "active space". Lirnax can show positive anemotaxis to gentle, l)resumably odor-free winds (31).
Olfactory stimuli are also important in locating food. Slugs
normally eat a variety of plants, including fungi (13). Removal of the optic tentacles containing the digitate ganglion
reduces the distance at which slugs detect the stinkhorn
(Phallus impudicus) from 120 cm to 20 cm (20). A similar result was obtained testing the response of A griolimax reticulatus
to potato before and after optic tentacle removal (21). Both
of these exl)erimelits indicated a residual olfactory sensitivity
after optic tentacle removal attributed to the smaller anterior tentacles. However, this l)henomenon was iiot al)l)arent
in my exl)eriments. The olfactory sensitivity and capacity
for directed locomotion via olfactory cues are also relevant
to ecological studies of slug distribution in relation to food

plant abundcance (32, 33).

The neurophysiological data obtained to date demonstrate
the olfactory function of the digitate ganglion in the slug. A
negative EOG in response to attractive plant odors has been
recorded from snail tentacles (28), whereas methanol and
ethanol produced a positive EOG. Based on data from frog
olfactory mucosa, Gesteland (34) suggests that substances
producing a negative EOG excite receptor cells and that substances producing a positive EOG inhibit receptor cells. It
will be very interesting to determine the odor response spectrum of single cells and so determine the presence of odor
generalists or odor specialists (35).
Input from the olfactory nerve enters the metacerebrum
(36) and ultimately impinges on pedal neurons that control
locomotion. Slugs move by producing waves of contraction
and elongation which move anteriorly over the foot (37). Although the production of these pedal waves has been reported to be independent of the central nervous system (38,
39), we have found that animals with sectioned pedal nerves
survive for weeks with no sign. of pedal wave activity in the
denervated region (Prior and Gelperin, unpublished observation). In the absence of any directive visual or olfactory stimuli, the circling locomotion which occurs (40) is probably due
to bilateral asymmetries in spontaneous pedal neuron outputs
(41).

FIG. 5. Al and B1 are recordings from olfactory nerve; A2 and

B2 are recordings from receptor surface. Al and A2 were recorded
simultaneously in response to 0.5 cc of amyl acetate vapor.
B1, B2 are the responses to 0.5 cc moist air. Calibration bar
applies to A and B and indicates 1 sec., 200 ,uV for Al, B1 and
400 ,AV for A2, B2.

The data presented here are very useful in designing experiments to probe plasticity of behavior in the slug. Olfaction,
taste, and vibration sensitivity (42) are the dominant senses
and learning paradigms must accommodate these facts.
Closely related snails have been trained to avoid previously
attractive plant odors by shocking them in the presence of the
odor (43). The aversive taste of quinine was effective in modifying the climbing behavior of Helix (44). Snails can also
learn to keep an optic tentacle retracted to avoid an aversive
stimulus (45). The recent demonstration of operant conditioning in Octopus (46) has further extended our knowledge of
the capabilities of molluscan brains.
Note Added in Proof. Recent experiments (47) demonstrate
that using olfactory cues, Limax can rapidly learn to avoid
new foodplants if aversive stimulation is paired with ingestion
of the new foodplant.
I thank D. Giesker for collecting some of these data. Supported
by N.S.F. Grant GB 20762.
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