Aggregation in the haematophagous bug Triatoma infestans: a novel assembling factor

Abstract. The response of Triatoma infestans (Klug) to odours left behind by walking insects was analysed. The insects aggregated on papers marked with putative chemical footprints in the absence of other assembling chemical signals. Contact of the receptor insect with the treated surface was necessary to elicit the response because the signal was not able to attract insects, only to arrest them on marked surfaces. These requirements contrast with other chemical aggregation signals of triatomines that attract the insect and involve olfactory responses. By washing the cuticle of the insects with hexane and impregnating papers with the extract, a response was evoked in test insects similar to that elicited by papers on which insects had walked. The biological role of this novel aggregation factor is discussed in relation to other assembling signals of different origin.     

The role of water vapour in the orientation behaviour of the blood-sucking bug Triatoma infestans (Hemiptera,Reduviidae)

The behavioural response to water vapour of the haematophagous bug Triatoma infestans was analysed. Dry or humid discrete sources at different temperatures were used as stimuli for insects walking on a locomotion compensator. Humidity significantly increased the tendency of these bugs to orientate towards thermal sources. Furthermore, humid sources at room temperature were attractive to T. infestans, but this effect was limited to short-range distances. On the other hand, dynamic sources, i.e. airstreams carrying different water vapour contents did not affect the spontaneous anemotactic behaviour of this species, neither in sign (positive) nor in intensity. The anemotactic behaviour was also not influenced by the physiological water balance state of the bugs. Results are discussed in relation to the cues released by living hosts of triatomine bugs and in relation to their responses to air-currents.     

Circadian control of photonegative sensitivity in the haematophagous bug Triatoma infestans

The negative phototactic response of Triatoma infestans (Hemiptera) was studied in an arena, half of which was kept dark and the other half illuminated with different light intensitites. For each intensity we measured the time the insects spent in the dark half, the time to reach the side opposite to that where they were released, and the number of passages through the middle line of the arena. T. infestans displayed a photonegative behaviour that was enhanced by high light intensities. Bugs maintained in 12:12 light-dark cycles responded differently to the same illumination levels when tested in their photophase and scotophase: sensitivity to light was higher during the latter. Bugs entrained to light-dark cycles, kept afterwards either in constant darkness or in constant light, and tested in their subjective night and day, showed the same responses as bugs from the light-dark group tested in their corresponding photophase and scotophase. Thus, phototactic sensitivity is under endogenous control. The behaviour shown by T. infestans may be understood as being composed of at least two different drives: an exploratory one, and a negative phototactic response that is under endogenous control and is particularly sensitive to light during the scotophase, when activity peaks occur. Accepted: 16 July 1998     

Visual lateral fixation and tracking in the haematophagous bug Triatoma infestans

Summary Triatoma bugs turn away from a vertical black stripe on the inner surface of a rotating or oscillating drum by keeping it in the lateral visual field at an angle relative to the long axis of the body. The value of depends on the angular velocity w of the drum. Below w=5° s^–1, increases with increasing w, and the stripe can lie to either side of the animal. Occasionally, the bugs switch between these two tracking modes. Above w=5° s^–1, remains nearly constant at about 120° and the stripe lags behind the animal. We call this lateral tracking. At velocities over 5° s^–1 the animals track the leading edge of a wide black stripe in the same manner as they track a narrow stripe. Below 5° s^–1 they walk towards the centre of the stripe (skototaxis). Objects moving towards the insect above the horizon are also fixated at an angle of about 120°. Lateral tracking is mediated mainly by the dorsal part of the visual field, as can be shown by occluding either the dorsal or ventral halves of the eyes. The walking speed of the bugs increases significantly during lateral tracking of an oscillating stripe compared with that during frontal fixation of a stationary one. We therefore interpret lateral tracking as an escape response.     

The central projection of cephalic mechanosensory axons in the haematophagous bug Triatoma infestans

The projections of mechanosensory hairs located on the dorsal and lateral head of the adult haematophagous bug Triatoma infestans were analyzed by means of cobalt filling. Axons run into the anterior and posterior tegumentary nerve and project through the brain to the ventral nerve cord. The fibres are small in diameter and run as a fascicle. Some branches run into suboesophageal and prothoracic centres; others run as far as to the mesothoracic ganglion. These sensory projections resemble that of wind-sensitive head hairs of the locust. The functional role of this sensory system in this species is discussed.     

Differential control of light-dark adaptation in the ocelli and compound eyes of Triatoma infestans

The adaptation to light of compound eyes in insects has been extensively documented and their adaptive role is well understood. Much less attention has been paid, however, to the control of ocelli sensitivity, a study which could help us to understand the functional role of these simple eyes. We analyzed the dynamic changes in the distribution of screening pigments which occur in the ocelli of the haematophagous bug, Triatoma infestans, when the insects are subjected either to light/dark cycles (LD), to constant darkness (DD) or constant light (LL). We then compared these changes with those occurring in the compound eyes of the same individuals and found that, while compound eyes are subject to the control of an endogenous circadian clock, the adaptation of the ocelli is entirely dependent on environmental illumination. In addition, we have observed that environmental temperature is not involved in the control of screening pigments in either ocelli or compound eyes as a direct stimulus, nor as a Zeitgeber. The existence of a differential control in the components of the dual visual system represents an adaptive advantage in the adjustment of visual sensitivity in insects exposed to quick changes in lighting conditions in their natural habitat. We discuss the implications of our findings with regards to the biology of triatomines and with respect to the general understanding the functional role of insect ocelli.     

Thermal preferences and limits of Triatoma brasiliensis in its natural environment - Field observations while host searching

The goal of this work was to explore the thermal relationship betweenforaging Triatoma brasiliensis and its natural habitat during thehottest season in the state of Ceará, Brazil. The thermal profiles were determinedusing infrared analysis. Although the daily temperature of rock surfaces varied in awide range, T. brasiliensis selected to walk through areas withtemperatures between 31.7-40.5ºC. The temperature of T. brasiliensisbody surface ranged from 32.8-34.4ºC, being higher in legs than the abdomen.A strong relationship was found between the temperature of the insect and thetemperature of rock crevices where they were hidden (r: 0.96, p < 0.05). Thespecies was active at full sunlight being a clear example of how the light-darkrhythm may be altered, even under predation risk. Our results strongly suggest athermal borderline for T. brasiliensis foraging activity near 40ºC.The simultaneous determination of insect body and rock temperatures here presentedare the only obtained in natural habitats for this or other triatomines.     

New structures for goat corrals to control peridomestic populations of Triatoma infestans (Hemiptera: Reduviidae) in the Gran Chaco of Argentina

Goat production is an important economic activity for rural communities in the Gran Chaco of Argentina. Goat corrals are important for the survival of peridomestic populations of Triatoma infestans. This study evaluated the impact of modifying the traditional structure of goat corrals on T. infestans populations and goat productivity in the region of Los Llanos (La Rioja). Thirty-nine experimental corrals were constructed and 57 traditional corrals were used as controls. We evaluated the infestations of the control and experimental corrals for five years following construction of the structures. The results showed that the new structures did not prevent the colonization, although it enhanced the detection of infestation at low densities of T. infestans. No significant difference was found in T. infestans population abundance between control and experimental corrals, probably because of the different detectability in the two types of structures, especially among the small nymphs. Although goat productivity average was higher in experimental than in control corrals, no significant difference was found because of high variability. The new structures can be used as a complement to promote the development of rural communities. Acceptability and adoption of the new corrals by the owners was high, as the enclosures offered better protection for the goats, increased growth of kids and facilitated herd handling.     

Daily rhythm of aggregation in the haematophagous bug Triatoma infestans (Heteroptera: Reduviidae)

Triatomine bugs show a temporal modulation of many activities. Here, we analyse the daily modulation of the aggregation behaviour of Triatoma infestans larvae and its chronobiological basis. In the laboratory, groups of six bugs were released over an experimental arena during six consecutive days, where their aggregation behaviour was quantified every hour. When submitted to a 12/12 h photoperiod (L/D), the larvae of T. infestans exhibited a cyclic pattern of aggregation with a 24 h period, evincing the existence of a daily rhythm of aggregation in this species. Bugs exhibited the maximum aggregation tendency at the end of the scotophase (7:00 h), moment in which they naturally search for refuges. The minimum aggregation (i.e. maximal dispersion) was observed during the last part of the photophase and beginning of the scotophase (15:00 to 1:00 h). This cyclic pattern disappeared when constant conditions of illumination (L/L) or darkness (D/D) were imposed to the bugs, suggesting the absence of an endogenous circadian control of this behaviour. Insects submitted to L/L and D/D photoperiods presented lower global levels of aggregation than those submitted to L/D conditions. The lack of an endogenous control and the relevance of light cycles as a synchronization signal are discussed as the temporal modulation of this behaviour might play an important role in the nocturnal habits of this species.     

Ammonia attracts the haematophagous bug Triatoma infestans : behavioural and neurophysiological data on nymphs

1) Nymphs of the haematophagous bug Triatoma infestans (Heteroptera: Reduviidae) are attracted to volatiles from their own faeces on a servosphere. 2) Biological substrates attractive to triatomines release NH₃: wetted triatomine faecal papers release NH₃ at 256 ppb NH₃ from a 60-g source and stale rabbit urine at 394 ppb from 200 ml. Ammonia released from aqueous NH₃ also attracts bugs at doses of 3 ppb and 17 ppb on the servosphere. 3) Bugs typically show negative anemotaxis in a stimulus-free air-stream on the servosphere. At onset of stimulation with ammonia from either biological substrates or aqueous NH₃ the bugs stop, move their antennae, turn and walk upwind, i.e. odour-mediated anemotaxis. 4) At lower NH₃ doses a latency in attraction is recorded, but this latency disappears when the relative humidity of the stimulus delivery air-stream is dropped from 90 to 35%. 5) Electrophysiological recordings from single olfactory sensilla on antennae of Triatoma nymphs reveal two different types of NH₃-excited receptors, both within grooved-peg sensilla. The responses of one of these receptor cells to NH₃ has been studied in detail and shows that the action potential discharge rate is dose-dependent over the range 2–200 ppb. 6) The amplitudes of electroantennograms recorded from Triatoma nymphs to NH₃ are dose dependent over the range 5–550 ppb. Accepted: 16 January 1997     

Degradation of serum proteins in Triatoma infestans.

It has been previously described (Perassi 1972, 1973)that no serum proteins from the animal on which the insects were fed can be detected in the insects' hemolymph of either male or female Triatoma infestans. In other words, the absorption of the proteins does not take place from the gut content to the hemolymph. Furthermore, according to the present investigation, the degradative processes of the gut content has revealed that, at least from an immunological point of view, an almost complete degradation of the rat serum proteins is carried out in the wide and slender midgut of T. infestans. These findings support the conclusion that in T. infestans the absorption of serum proteins from the gut to the hemolymph does not occur unless the proteins have been degradated previously.     

The role of antennae in the thermopreference and biting response of haematophagous bugs

Insects sense thermal cues mainly through thermoreceptors located in the antenna. To analyse the impact of antennectomy on the thermal behaviour of the haematophagous bug Triatomainfestans, we studied the distribution of intact and antennectomised bugs in an experimental arena where a temperature gradient was established, as well as the biting response of insects with and without antennae to objects at the temperature of a potential host. Antennectomy did not abolish thermopreference, but modified the temperature at which the insects preferred to stay. In the arena, antennectomised insects chose to remain at a higher temperature (ca. 3 °C higher in unfed bugs), and exhibited a larger dispersion around that preferred temperature, than intact bugs. In addition, ablated insects temporarily lost their ability to bite an object at the temperature of a potential host, but that ability was gradually recovered after the fifth day post-antennectomy. Results presented here show that thermoreceptors other than those located on the antennae can also guide thermal behaviours. We conclude that the function of antennal thermoreceptors can be taken over by other receptors located in different regions of the body. Those receptors have a different sensitivity and confer the insects with a different responsiveness.     

Population dynamics and control of Triatoma infestans

Analysis of field populations of Triatoma infestans (Hemiptera: Reduviidae), after a 3-year study, shows that population growth rate is affected by both density-dependent and density-independent mortality. Although an equilibrium exists, apparently as a consequence of a density dependent-mechanism, population density fluctuates throughout the year because of the effect of monthly mean minimum temperature as a density-independent source of mortality. Simulation studies based on Moran curves shows that high population densities have an approximately constant extinction probability (around 0.20), independently of the season the population starts growing. However, at very low population densities, the extinction probability depends strongly on the season when the population begins to grow. Very low density populations beginning in winter or autumn have the highest extinction probability. The outcome of the simulation studies coincides with results observed in field populations affected by insecticide application at different seasons.     

The biting rate of Triatoma infestans in Argentina

1. The daily proportion of fed individuals in a population of the reduviid bug Triatoma infestans (Klug), maintained under natural climatic conditions in experimental chicken houses in central Argentina, was estimated from the proportion of bugs retaining colourless urine in the rectum. 2. From the estimates of feeding frequency throughout a 1 year period, it was shown that temperature has a dominant effect on biting rate, but density-dependent effects became apparent during the warmest months. 3. These and other data on the determinants of blood consumption by T. infestans, were incorporated into a detailed hypothesis of density regulation in this species.     

Population dynamics of Triatoma infestans under natural climatic conditions in the Argentine Chaco

Using experimental chicken houses at a site in central Argentina where the bug Triatoma infestans (Klug) is endemic, nine populations of this vector of Chagas disease were monitored during a 34-month period. Bug populations with four chickens as hosts were consistently larger than those with two chickens as hosts. Age structure of the bug population followed a similar pattern irrespective of the initial age structure. Egg to adult mortality was consistently around 98.5% and there was no consistent evidence for density-dependent mortality. There was some evidence for density dependence in fecundity and recruitment rates, but these were heavily constrained by low temperatures during the winter months. Nymphal development rates correlated most strongly with mean minimum temperatures rather than with mean maximum temperatures. We conclude that vector control using insecticides against this species would be most effective at the onset of winter, when recovery of any surviving populations would be inhibited by low temperatures.     

Population morphometric analysis of the tropicopolitan bug Triatoma rubrofasciata and relationships with Old World species of Triatoma: evidence of New World ancestry

Quantitative analysis of morphological characters of the head was used to reconstruct the evolutionary history of the tropicopolitan bug Triatoma rubrofasciata (De Geer) (Hemiptera: Reduviidae) and seven species of Old World Triatoma. Multivariate analysis demonstrates that T. rubrofasciata and the Old World species have a high degree of similarity with Nearctic Triatoma species, particularly T. rubida (Uhler). We interpret this to imply a common ancestry for these groups. Dissemination of T. rubrofasciata and subsequent derivation of the Old World species of Triatoma is deduced to have occurred over a period of not more than 350 years.     

Novel polymerase chain reaction-restriction fragment length polymorphism assay to determine internal transcribed spacer-2 group in the Chagas disease vector,Triatoma dimidiata (Latreille, 1811)

Triatoma dimidiata is the most important Chagas disease insect vector in Central America as this species is primarily responsible for Trypanosoma cruzi transmission to humans, the protozoan parasite that causes Chagas disease. T. dimidiata sensu lato is a genetically diverse assemblage of taxa and effective vector control requires a clear understanding of the geographic distribution and epidemiological importance of its taxa. The nuclear ribosomal internal transcribed spacer 2 (ITS-2) is frequently used to infer the systematics of triatomines. However, oftentimes amplification and sequencing of ITS-2 fails, likely due to both the large polymerase chain reaction (PCR) product and polymerase slippage near the 5'' end. To overcome these challenges we have designed new primers that amplify only the 3''-most 200 base pairs of ITS-2. This region distinguishes the ITS-2 group for 100% of known T. dimidiata haplotypes. Furthermore, we have developed a PCR-restriction fragment length polymorphism (RFLP) approach to determine the ITS-2 group, greatly reducing, but not eliminating, the number of amplified products that need to be sequenced. Although there are limitations with this new PCR-RFLP approach, its use will help with understanding the geographic distribution of T. dimidiata taxa and can facilitate other studies characterising the taxa, e.g. their ecology, evolution and epidemiological importance, thus improving vector control.     

Determinants of the domiciliary density of Triatoma infestans, vector of Chagas disease

In two heavily infested rural villages of Santiago del Estero, Argentina, where no indoor-spraying with residual insecticides had ever been carried out by official control services, we studied the influence of roof and wall structure, domestic use of insecticide, family size and the number of domestic dogs, on the domiciliary density of Triatoma infestans (Klug). Bug density was significantly associated with (1) the interaction between insecticide use and type of roof, (2) the structure of indoor walls, (3) the number of dogs sharing sleeping areas of people (room-mate dogs), and (4) the number of people plus room-mate dogs, but not with just the number of people resident in the house. The interaction between insecticide use and a roof made of 'simbol', a locally available grass (Pennisetum sp.), also reflected a younger age structure of domestic bug populations. In infested houses, the density of bugs infected with Trypanosoma cruzi Chagas was significantly correlated with overall bug density. Our data suggest that the application of environmental management measures by the affected people, such as plastering of walls and modification of roofs, coupled with keeping dogs away from bedrooms and application of insecticides, should limit the domestic population density of T. infestans and thus reduce the transmission of T. cruzi to people.     

Repellency of deet to nymphs of Triatoma infestans

The repellency of N,N-diethyl-3-methylbenzamide (deet) to Triatoma infestans (Klug) (Hemiptera: Reduviidae) was evaluated using third-instar nymphs and a video tracking technique. Three experimental designs were used: (a) the test arena floor was divided into two halves, only one of which was treated with deet; (b) the arena floor was divided into an inner circle and an outer ring, only the latter treated with deet; (c) half of the test arena was covered by a filter paper roof treated with deet (out of reach of the nymphs). In all three types of experiment, a repellent effect was demonstrated proportional to the dose of deet. When a host (pigeon) was shielded by deet-treated gauze, the rate of blood-feeding by the nymphs was inhibited. Topical pre-treatment of the nymphs with N-ethylmaleimide, to block chemoreception, inhibited the repellency.