The Ant Who Cried Wolf? Short-Term Repeated Exposure to Alarm Pheromone Reduces Behavioral Response in Argentine Ants.

In this study we test whether Argentine ants (Linepithema humile) progressively reduce their response to a salient stimulus (alarm pheromone) with increased exposure over time. First, we used a two-chamber olfactometer to demonstrate three focal behaviors of Argentine ants that indicate an alarmed state in response to conspecific alarm pheromone and pure synthetic iridomyrmecin (a dominant component of L. humile alarm pheromone). We then measured how these behaviors changed after repeated exposure to conspecific alarm pheromone from live ants. In addition, we investigate whether there is a difference in the ants' behavioral response after "short" (3 min) versus "long" (6 min) intervals between treatments. Our results show that Argentine ants do exhibit reduced responses to their own alarm pheromone, temporarily ceasing their response to it after four or five exposures, and this pattern holds whether exposure is repeated after "short" or "long" intervals. We suggest alarm pheromones may be perceived as false alarms unless threatening stimuli warrant a continued state of alarm. These results should be kept in mind while developing pheromone-based integrated pest management strategies.

The nature of alarm communication in Constrictotermes cyphergaster (Blattodea: Termitoidea: Termitidae): the integration of chemical and vibroacoustic signals.

Alarm signalling is of paramount importance to communication in all social insects. In termites, vibroacoustic and chemical alarm signalling are bound to operate synergistically but have never been studied simultaneously in a single species. Here, we inspected the functional significance of both communication channels in Constrictotermes cyphergaster (Termitidae: Nasutitermitinae), confirming the hypothesis that these are not exclusive, but rather complementary processes. In natural situations, the alarm predominantly attracts soldiers, which actively search for the source of a disturbance. Laboratory testing revealed that the frontal gland of soldiers produces a rich mixture of terpenoid compounds including an alarm pheromone. Extensive testing led to identification of the alarm pheromone being composed of abundant monoterpene hydrocarbons (1S)-α-pinene and myrcene, along with a minor component, (E)-ß-ocimene. The vibratory alarm signalling consists of vibratory movements evidenced as bursts; a series of beats produced predominantly by soldiers. Exposing termite groups to various mixtures containing the alarm pheromone (crushed soldier heads, frontal gland extracts, mixture of all monoterpenes, and the alarm pheromone mixture made of standards) resulted in significantly higher activity in the tested groups and also increased intensity of the vibratory alarm communication, with the responses clearly dose-dependent. Lower doses of the pheromone provoked higher numbers of vibratory signals compared to higher doses. Higher doses induced long-term running of all termites without stops necessary to perform vibratory behaviour. Surprisingly, even crushed worker heads led to low (but significant) increases in the alarm responses, suggesting that other unknown compound in the worker's head is perceived and answered by termites. Our results demonstrate the existence of different alarm levels in termites, with lower levels being communicated through vibratory signals, and higher levels causing general alarm or retreat being communicated through the alarm pheromone.

An alarm pheromone increases the responsivity of amygdaline-hippocampal neurons / Una feromona de alarma incrementa la responsividad de neuronas amigdalino-hipocampales

The capability to perceive and emit alarm substances, such as 2-heptanone, makes animals aware of the presence of danger, leading to some strategies directed towards survival. Strategies of survival involve emotional memory which is processed by deep temporal lobe structures, such as amygdaloid complex and hippocampus. In the Wistar rat, 2-heptanone produces anxiety-like behavior and an increased firing rate of basal amygdaline neurons. However, it is unknown whether 2-heptanone modifies the responsivity of medial amygdaline-hippocampal connection. Therefore, we placed a group (n=10) of Wistar rats in a plexiglass cage impregnated with 2-heptanone. Rats from control group (n=10) were introduced into a similar clean cage. Twenty four hours later we obtained single-unit extracellular recordings from the hippocampus (CA1-CA3) neurons identified by their connection to medial amygdala. Although the basal neuronal firing rate was similar between groups, first order interval distribution histogram analysis showed that 2-heptanone produced shorter intervals of firing rate. Peristimulus histograms indicated that: i) the amygdaline stimulation produces an increased firing rate in hippocampal neurons; and ii) this response is increased and enlarged on the 2-heptanone group. Since a single exposure to an alarm pheromone seems to facilitate the amygdala-hippocampal connection, results suggest the initial formation of contextual memories related with fear.

La percepción olfatoria de feromonas de alarma, como la 2-heptanona, promueve ciertas estrategias de supervivencia con la participación de la memoria emocional, integrada en estructuras del lóbulo temporal, como la amígdala y el hipocampo. En la rata Wistar, la olfacción de 2-heptanona genera conductas sugerentes de ansiedad y un incremento de la tasa de disparo neuronal del núcleo basal de la amígdala. Sin embargo, no se conoce si la 2-heptanona modifica la responsividad de la conexión amígdala medial-hipocampo. Un grupo de ratas Wistar (n=10) fue colocado dentro de una caja de acrílico impregnada con 2-heptanona; el grupo control (n=10) fue introducido en una caja limpia. Veinticuatro horas después se obtuvo el registro unitario extracelular de neuronas del hipocampo (CA1-CA3) identificadas por su conexión con la amígdala medial. Aunque la tasa de disparo basal fue similar entre los grupos experimentales, el histograma de distribución de intervalos de primer orden indicó un predominio de intervalos de breve duración en el grupo 2-heptanona. Los histogramas periestímulo indicaron que: i) las neuronas hipocampales responden con un incremento en la tasa de disparo neuronal ante la estimulación amigdalina; ii) la respuesta es de mayor magnitud y duración en el grupo previamente expuesto a 2-heptanona. Dado que una sola exposición a una feromona de alarma facilita la conexión amígdala medial-hipocampo, los resultados sugieren la formación inicial de una memoria contextual relacionada con el miedo.

The main component of an alarm pheromone of kissing bugs plays multiple roles in the cognitive modulation of the escape response.

Innate responses in animals can be modulated by experience. Disturbed adults of the triatomine bug Triatoma infestans release an alarm pheromone (AP) that elicits an escape response in conspecific larvae. The main component of this AP, the isobutyric acid (IsoAc), alone has already shown to generate an escape response in this species. However, not much is known about the modulation of this behavior by non-associative and associative cognitive processes. We present here evidences of the cognitive capacities of T. infestans larvae in an escape context under different conditioning paradigms, including IsoAc in different roles. We show that: (1) the duration of a pre-exposure to IsoAc plays a main role in determining the type of non-associative learning expressed: short time pre-exposures elicit a sensitization while a longer pre-exposure time triggers a switch from repellence to attractiveness; (2) a simple pre-exposure event is enough to modulate the escape response of larvae to the AP and to its main component: IsoAc; (3) IsoAc and the AP are perceived as different chemical entities; (4) an association between IsoAc and an aversive stimulus can be created under a classical conditioning paradigm; (5) an association between IsoAc and a self-action can be generated under an operant conditioning. These results evince that IsoAc can attain multiple and different cognitive roles in the modulation of the escape response of triatomines and show how cognitive processes can modulate a key behavior for surviving, as it is the escaping response in presence of a potential danger in insects.

Chemical communication in isoptera / Comunicação química em isoptera

The semiochemicals produced by termites have been little studied compared to those of ants and bees. Among the limiting factors are the cryptic habits of termites, together with the difficulty in maintaining many species in the laboratory. The semiochemicals of Isoptera include trail, sex and alarm pheromones and a complex mixture of substances responsible for the recognition of nestmates. Although little is known about the semiochemicals of termites, available data indicate a strategy of pheromonal parsimony in the chemistry communication of these insects, i.e., the same compound is sometimes secreted by different glands, different species and for different functions.

Os semioquímicos dos cupins têm sido pouco estudados quando comparados aos de formigas e abelhas. Um dos fatores limitantes é o hábito críptico dos térmitas, aliado à dificuldade de manutenção de muitas espécies em laboratório. Entre os semioquímicos de Isoptera destacam-se os feromônios sexuais, de trilha, de alarme e uma mistura complexa de substâncias responsável pelo reconhecimento dos companheiros de ninho. Apesar de os semioquímicos dos cupins serem pouco conhecidos, os dados disponíveis indicam uma estratégia de parcimônia feromonal na comunicação química desses insetos, ou seja, algumas vezes o mesmo composto é secretado por diferentes glândulas, diferentes espécies e para diferentes funções.