Ants Can Learn to Forage on One-Way Trails

The trails formed by many ant species between nest and food source are two-way roads on which outgoing and returning workers meet and touch each other all along. The way to get back home, after grasping a food load, is to take the same route on which they have arrived from the nest. In many species such trails are chemically marked by pheromones providing orientation cues for the ants to find their way. Other species rely on their vision and use landmarks as cues. We have developed a method to stop foraging ants from shuttling on two-way trails. The only way to forage is to take two separate roads, as they cannot go back on their steps after arriving at the food or at the nest. The condition qualifies as a problem because all their orientation cues-chemical, visual or any other - are disrupted, as all of them cannot but lead the ants back to the route on which they arrived. We have found that workers of the leaf-cutting ant Atta sexdens rubropilosa can solve the problem. They could not only find the alternative way, but also used the unidirectional traffic system to forage effectively. We suggest that their ability is an evolutionary consequence of the need to deal with environmental irregularities that cannot be negotiated by means of excessively stereotyped behavior, and that it is but an example of a widespread phenomenon. We also suggest that our method can be adapted to other species, invertebrate and vertebrate, in the study of orientation, memory, perception, learning and communication.

Harvest-ironman: heavy armature, and not its defensive secretions, protects a harvestman against a spider

Natural selection has caused prey species to evolve distinct defensive mechanisms. One of such mechanisms was the evolution of noxious or distasteful chemicals, which have appeared independently in a number of vertebrates and invertebrates. In detailed analyses of arthropod behaviour, scent gland secretions have consistently been shown to be responsible for repelling specific predators. Because using such chemicals is costly, animals with alternative cheaper defences are expected not to release such secretions when alternative options exist. In this study, we sought to determine the defensive mechanisms of the harvestman Discocyrtus invalidus, a heavy bodied species that bears a pair of repugnatorial glands. The spider Enoploctenus cyclothorax was used as the predator, and the cricket Gryllus sp. was used as a control. In a first set of experiments, the harvestmen were preyed upon significantly less than the crickets. In two other experiments, we found that harvestmen did not use their scent gland secretions to deter the predator. Moreover, results of a fourth experiment revealed that these spiders are not repelled by defensive secretions. Discocyrtus invalidus has a thick cuticle on the entire body: scanning electron micrographs revealed that only the mouth, the articulations of appendages and the tips of the legs are not covered by a hard integument. In a fifth experiment, we found that these spiders had difficulty piercing the harvestmen body. This is the first experimental evidence that a chemically defended arachnid does not use its scent gland secretions to repel a much larger predator but instead relies on its heavily built body. (c) 2010 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

A novel multiobjective optimization algorithm based on bacterial chemotaxis

In this article a novel algorithm based on the chemotaxis process of Echerichia coil is developed to solve multiobjective optimization problems. The algorithm uses fast nondominated sorting procedure, communication between the colony members and a simple chemotactical strategy to change the bacterial positions in order to explore the search space to find several optimal solutions. The proposed algorithm is validated using 11 benchmark problems and implementing three different performance measures to compare its performance with the NSGA-II genetic algorithm and with the particle swarm-based algorithm NSPSO. (C) 2009 Elsevier Ltd. All rights reserved.

Artificial neural networks analysis of Sao Paulo subway tunnel settlement data

Artificial neural networks have been used to analyze a number of engineering problems, including settlement caused by different tunneling methods in various types of ground mass. This paper focuses on settlement over shotcrete- supported tunnels on Sao Paulo subway line 2 (West Extension) that were excavated in Tertiary sediments using the sequential excavation method. The adjusted network is a good tool for predicting settlement above new tunnels to be excavated in similar conditions. The influence of network training parameters on the quality of results is also discussed. (C) 2007 Elsevier Ltd. All rights reserved.

Influence of physical and geometrical parameters on three-dimensional load transfer mechanism at tunnel face

Three-dimensional discretizations used in numerical analyses of tunnel construction normally include excavation step lengths much shorter than tunnel cross-section dimensions. Simulations have usually worked around this problem by using excavation steps that are much larger than the actual physical steps used in a real tunnel excavation. In contrast, the analyses performed in this study were based on finely discretized meshes capable of reproducing the excavation lengths actually used in tunnels, and the results obtained for internal forces are up to 100% greater than those found in other analyses available in the literature. Whereas most reports conclude that internal forces depend on support delay length alone, this study shows that geometric path dependency (reflected by excavation round length) is very strong, even considering linear elasticity. Moreover, many other solutions found in the literature have also neglected the importance of the relative stiffness between the ground mass and support structure, probably owing to the relatively coarse meshes used in these studies. The analyses presented here show that relative stiffness may account for internal force discrepancies in the order of 60%. A dimensionless expression that takes all these parameters into account is presented as a good approximation for the load transfer mechanism at the tunnel face.

Sites of Defoliation by Atta sexdens rubropilosa (Hymenoptera, Formicidae) in Artificial Plants

This study aimed to achieve a better understanding about the foraging behavior of leaf-cutter ant (Atta sexdens rubropilosa Forel) workers with respect to defoliation sites in plants. To accomplish that, artificial plants 70 cm in height were prepared and divided into four levels (heights), having natural plant leaves attached to them. Evaluations during the bioassays included the number of leaves dropped by the ants, as well as the percentage of plant mass removed. In all replicates, it became evident that the most exploited plant site is the apical region, which significantly differed from other plant levels.

Nestmate recognition in a stingless bee: does the similarity of chemical cues determine guard acceptance?

The ability to discriminate nestmates from non-nestmates is critical to the maintenance of the integrity of social insect colonies. Guard workers compare the chemical cues of an incoming individual with their internal template to determine whether the entrant belongs to their colony. In contrast to honeybees, Apis mellifera, stingless bees have singly mated queens and, therefore, are expected to have a higher chemical homogeneity in their colonies. We tested whether aggressive behaviour of Frieseomelitta varia guards towards nestmate and non-nestmate foragers reflects chemical similarities and dissimilarities, respectively, of cuticular hydrocarbon profiles. We also introduced individuals of Lestrimelitta limao, an obligatory robber species, to test the ability of guards to react effectively to intruders from other taxa. We verified that foraging nestmates were almost invariably accepted, while heterospecific and conspecific non-nestmates were rejected at relatively high rates. However, non-nestmate individuals with higher chemical profile similarity were likely to be accepted by guards. We conclude that guards compare the chemical cuticular blend of incoming individuals and make acceptance decisions according to the similarity of the compounds between the colonies. (c) 2007 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Olfactory eavesdropping between two competing stingless bee species

Foragers can improve search efficiency, and ultimately fitness, by using social information: cues and signals produced by other animals that indicate food location or quality. Social information use has been well studied in predator-prey systems, but its functioning within a trophic level remains poorly understood. Eavesdropping, use of signals by unintended recipients, is of particular interest because eavesdroppers may exert selective pressure on signaling systems. We provide the most complete study to date of eavesdropping between two competing social insect species by determining the glandular source and composition of a recruitment pheromone, and by examining reciprocal heterospecific responses to this signal. We tested eavesdropping between Trigona hyalinata and Trigona spinipes, two stingless bee species that compete for floral resources, exhibit a clear dominance hierarchy and recruit nestmates to high-quality food sources via pheromone trails. Gas chromatography-mass spectrometry of T. hyalinata recruitment pheromone revealed six carboxylic esters, the most common of which is octyl octanoate, the major component of T. spinipes recruitment pheromone. We demonstrate heterospecific detection of recruitment pheromones, which can influence heterospecific and conspecific scout orientation. Unexpectedly, the dominant T. hyalinata avoided T. spinipes pheromone in preference tests, while the subordinate T. spinipes showed neither attraction to nor avoidance of T. hyalinata pheromone. We suggest that stingless bees may seek to avoid conflict through their eavesdropping behavior, incorporating expected costs associated with a choice into the decision-making process.

Identification of a juvenile hormone esterase-like gene in the honey bee, Apis mellifera L. - Expression analysis and functional assays

Tight control over circulating juvenile hormone (JH) levels is of prime importance in an insect`s life cycle. Consequently, enzymes involved in JH metabolism, especially juvenile hormone esterases (JHEs), play major roles during metamorphosis and reproduction. In the highly eusocial Hymenoptera, JH has been co-opted into additional functions, primarily in the development of the queen and worker castes and in age-related behavioral development of workers. Within a set of 21 carboxylesterases predicted in the honey bee genome we identified one gene (Amjhe-like) that contained the main functional motifs of insect JHEs. Its transcript levels during larval development showed a maximum at the switch from feeding to spinning behavior, coinciding with a JH titer minimum. In adult workers, the highest levels were observed in nurse bees, where a low JH titer is required to prevent the switch to foraging. Functional assays showed that Amjhe-like expression is induced by JH-III and suppressed by 20-hydroxyecdysone. RNAi-mediated silencing of Amjhe-like gene function resulted in a six-fold increase in the JH titer in adult worker bees. The temporal profile of Amjhe-like expression in larval and adult workers, the pattern of hormonal regulation and the knockdown phenotype are consistent with the function of this gene as an authentic JHE. (C) 2008 Elsevier Inc. All rights reserved.

Worker demography in a large-colony, swarm-founding wasp

Neotropical swarm-founding wasps build nests enclosed in a covering envelope, which makes it difficult to count individual births and deaths. Thus, knowledge of worker demography is very limited for swarm-founding species compared with that for independent-founding species. In this study, we explored the worker demography of the swarm-founding wasp Polybia paulista, the colony size of which usually exceeds several thousand adults. We considered each wasp colony as an open-population and estimated the survival probability, recruitment rate, and population size of workers using the developments of the Cormack-Jolly-Seber model. We found that capture probability varied considerably among the workers, probably due to age polyethism and/or task specialization. The daily survival rate of workers was high (around 0.97) throughout the season and was not related to the phase of colony development. On the other hand, the recruitment rate ranged from 0 to 0.37, suggesting that worker production was substantially less important than worker survival in determining worker population fluctuations. When we compared survival rates among worker groups of one colony, the mean daily survival rate was lower for founding workers than for progeny workers and tended to be higher in progeny workers that emerged in winter. These differences in survivorship patterns among worker cohorts would be related to worker foraging activity and/or level of parasitism.

Developmental characterization, function and regulation of a Laccase2 encoding gene in the honey bee, Apis mellifera (Hymenoptera, Apinae)

In insects, exoskeleton (cuticle) formation at each molt cycle includes complex biochemical pathways wherein the laccase enzymes (EC 1.10.3.2) may have a key role. We identified an Amlac2 gene that encodes a laccase2 in the honey bee, Apis mellifera, and investigated its function in exoskeleton differentiation. The Amlac2 gene consists of nine exons resulting in an ORE of 2193 nucleotides. The deduced translation product is a 731 amino acid protein of 81.5 kDa and a pl of 6.05. Amlac2 is highly expressed in the integument of pharate adults, and the expression precedes the onset of cuticle pigmentation and the intensification of sclerotization. In accordance with the temporal sequence of exoskeleton differentiation from anterior to posterior direction, the levels of Amlac2 transcript increase earlier in the thoracic than in the abdominal integument. The gene expression lasts even after the bees emerge from brood cells and begin activities in the nest, but declines after the transition to foraging stage, suggesting that maturation of the exoskeleton is completed at this stage. Post-transcriptional knockdown of Amlac2 gene expression resulted in structural abnormalities in the exoskeleton and drastically affected adult eclosion. By setting a ligature between the thorax and abdomen of early pupae we could delay the increase in hemolymph ecdysteroid levels in the abdomen. This severely impaired the increase in Amlac2 transcript levels and also the differentiation of the abdominal exoskeleton. Taken together, these results indicate that Amlac2 expression is controlled by ecdysteroids and has a critical role in the differentiation of the adult exoskeleton of honey bees. (C) 2010 Elsevier Ltd. All rights reserved.

Reproductive biology of Cyrtopodium polyphyllum (Orchidaceae): a Cyrtopodiinae pollinated by deceit

The genus Cyrtopodium comprises about 42 species distributed from southern Florida to northern Argentina. Cyrtopodium polyphyllum occurs on rocks or in sandy soils, in restinga vegetation along the Brazilian coast. It flowers during the wet season and its inflorescences produce a high number of resupinate yellow flowers. Cyrtopodium polyphyllum offers no rewards to its pollinators, but mimics the yellow, reward-producing flowers of nearby growing Stigmaphyllon arenicola (oil) and Crotalaria vitellina (nectar) individuals. Several species of bee visit flowers of C. polyphyllum, but only two species of Centris (Centris tarsata and Centris labrosa) act as pollinators. Visits to flowers of C. polyphyllum were scarce and, as a consequence, low-fruit set was recorded under natural conditions. Such low-fruit production contrasts with the number of fruits each plant bears after manual pollination, suggesting deficient pollen transfer among plants. C. polyphyllum is self-compatible and has a high-fruit set in both manual self- and cross-pollinated flowers. Furthermore, fruits (2%) are formed by self-pollination assisted by rain. This facultative self-pollination mechanism is an important strategy to provide reproductive assurance to C. polyphyllum as rainfall restricts the foraging activity of its pollinating bees. Fruits derived from treatments and under natural conditions had a similar high rate of potentially viable seed. Moreover, these seeds had a low polyembryony rate, which did not exceed 5%. C. polyphyllum acts by deceit involving optical signals and exploits other yellow-flowered species within its habitat by attracting their pollinators. The low capsule production under natural conditions was expected, but its reproductive success is assured through self-pollination by rain and high seed viability.

The nymph of Tortopus harrisi Traver (Ephemeroptera: Polymitarcyidae)

Polymitarcyidae is a family of burrowing mayflies (Ephemeroptera: Ephemeroidea) distributed throughout the world but with highest diversity in the Neotropics. Tortopus Needham & Murphy, with a Panamerican distribution, is known from twelve species described in the adult stage. Nymphs are only known for three species: T. puella (Pictet), T. obscuripennis Dominguez and T. sarae Dominguez, and present a rather homogeneous morphology (Molineri 2008). They were firstly described for T. puella by Scott et al. (1959) and later Molineri (2008) described the other two. Both studies reported that these species burrow U-shaped tunnels in clay banks of rivers and streams, thus preventing them from being sampled in most limnological studies (that use surbers, drags, or drift nets). The aim of the present contribution is to describe and illustrate the previously unknown nymph of Tortopus harrisi Traver that shows important anatomical differences with the other nymphs known in the genus. This morphological differentiation suggests a different habitat use by these nymphs, sampled with drag and surber samplers in sandy substrate. New locality records are given for T. harrisi in Brazil. The nymphs are preserved in alcohol, mouthparts, legs and genital rudiments were mounted in microscope slides with Canada Balsam. Drawings were made with a camera lucida attached to a stereo microscope. The material is deposited in CUIC (Cornell University Insect Collection, Ithaca, NY), IML (Instituto Miguel Lillo, Tucuman) and in MZSP (Museu de Zoologia da Universidade de Sao Paulo, Sao Paulo). Catalogs and bibliography were consulted at Ephemeroptera Galactica (Hubbard 2009).

RNAi-mediated silencing of vitellogenin gene function turns honeybee (Apis mellifera) workers into extremely precocious foragers

The switch from within-hive activities to foraging behavior is a major transition in the life cycle of a honeybee (Apis mellifera) worker. A prominent regulatory role in this switch has long been attributed to juvenile hormone (JH), but recent evidence also points to the yolk precursor protein vitellogenin as a major player in behavioral development. In the present study, we injected vitellogenin double-stranded RNA (dsVg) into newly emerged worker bees of Africanized genetic origin and introduced them together with controls into observation hives to record flight behavior. RNA interference-mediated silencing of vitellogenin gene function shifted the onset of long-duration flights (> 10 min) to earlier in life (by 3-4 days) when compared with sham and untreated control bees. In fact, dsVg bees were observed conducting such flights extremely precociously, when only 3 days old. Short-duration flights (< 10 min), which bees usually perform for orientation and cleaning, were not affected. Additionally, we found that the JH titer in dsVg bees collected after 7 days was not significantly different from the controls. The finding that depletion of the vitellogenin titer can drive young bees to become extremely precocious foragers could imply that vitellogenin is the primary switch signal. At this young age, downregulation of vitellogenin gene activity apparently had little effect on the JH titer. As this unexpected finding stands in contrast with previous results on the vitellogenin/JH interaction at a later age, when bees normally become foragers, we propose a three-step sequence in the constellation of physiological parameters underlying behavioral development.

Insulin-like peptide genes in honey bee fat body respond differently to manipulation of social behavioral physiology

Nutrient sensitive insulin-like peptides (ILPs) have profound effects on invertebrate metabolism, nutrient storage, fertility and aging. Many insects transcribe ILPs in specialized neurosecretory cells at changing levels correlated with life history. However, the major site of insect metabolism and nutrient storage is not the brain, but rather the fat body, where functions of ILP expression are rarely studied and poorly understood. Fat body is analogous to mammalian liver and adipose tissue, with nutrient stores that often correlate with behavior. We used the honey bee (Apis mellifera), an insect with complex behavior, to test whether ILP genes in fat body respond to experimentally induced changes of behavioral physiology. Honey bee fat body influences endocrine state and behavior by secreting the yolk protein precursor vitellogenin (Vg), which suppresses lipophilic juvenile hormone and social foraging behavior. In a two-factorial experiment, we used RNA interference (RNAi)-mediated vg gene knockdown and amino acid nutrient enrichment of hemolymph (blood) to perturb this regulatory module. We document factor-specific changes in fat body ilp1 and ilp2 mRNA, the bee`s ILP-encoding genes, and confirm that our protocol affects social behavior. We show that ilp1 and ilp2 are regulated independently and differently and diverge in their specific expression-localization between fat body oenocyte and trophocyte cells. Insect ilp functions may be better understood by broadening research to account for expression in fat body and not only brain.