A programme of self-steered turns as a humidity response in Tenebrio, and the problem of categorizing spatial manoeuvres

ABSTRACT. Dry-acclimatized mealworm beetles, Tenebrio molitor L. (Coleoptera: Tenebrionidae), were exposed to a rapid change from dry to moist air in an arena with vertical air flow providing spatially uniform humidity, thus precluding orientation along humidity gradients. Subsequent angles of turn were significantly clustered around 90 and 180, and took the beetles back into the region from which they had come. The possibility of geomagnetic orientation was ruled out experimentally and in the apparent absence of other external directional cues it is concluded that the turn sequences were idiothetically controlled and pre-programmed. This and other cases of 'longitudinal' klinotaxis are discussed in the wider context of kineses and taxes and the currently-questioned value of these categories.     

Hygro- and thermoreceptive tarsal organ in the spider Cupiennius salei

Extracellular recordings were made from moist cells, dry cells and warm cells in the tip pore sensilla of the spider tarsal organ. Stimulation consisted of a rapid shift from an adapting air stream to another one at different levels of partial pressure of water vapor or of temperature. The moist and the dry cells respond antagonistically to sudden changes in humidity. Both hygroreceptors are unusual in being excited in a synergistic manner by pungent vapors of very volatile, polar substances. Presumably, the hygrosensitivity is superimposed on basically chemosensitive receptors. A moist cell at average differential sensitivity is able to discriminate two successive upward steps in humidity when they differ by 11% relative humidity. For a single dry cell, the difference required for a correct discrimination between two downward humidity steps is 10% relative humidity. The moist and the dry cells are unique in that they occur in combination with warm cells. A single warm cell at average differential sensitivity is able to resolve differences in warming steps down to 0.4°C.Abbreviations HR relative humidity - T temperature     

Movement behaviour of the pine weevil Hylobius abietis in relation to soil type: an arena experiment

Pine weevil, Hylobius abietis (L.) (Coleoptera: Curculionidae), movement behaviour on different substrates was studied in an indoor arena using a video camera and digital image processing technique. We analysed individual variation in movement characteristics, i.e. turning angles, movement directions, and distance moved per unit time on the bare and level arena surface which consisted of mineral soil (sand) and/or humus sections in various spatial configurations. Pine weevils on humus did not turn back when they came to the border with the sand. However, most individuals moved faster on sand than on humus. Thus, the results suggest that interactions between substrate differences and individual movement behaviour may to some extent explain why pine weevils have been observed to feed less frequently on coniferous seedlings planted on mineral soil than on those planted on humus.     

The Evaporative Function of Cockroach Hygroreceptors

Insect hygroreceptors associate as antagonistic pairs of a moist cell and a dry cell together with a cold cell in small cuticular sensilla on the antennae. The mechanisms by which the atmospheric humidity stimulates the hygroreceptive cells remain elusive. Three models for humidity transduction have been proposed in which hygroreceptors operate either as mechanical hygrometers, evaporation detectors or psychrometers. Mechanical hygrometers are assumed to respond to the relative humidity, evaporation detectors to the saturation deficit and psychrometers to the temperature depression (the difference between wet-bulb and dry-bulb temperatures). The models refer to different ways of expressing humidity. This also means, however, that at different temperatures these different types of hygroreceptors indicate very different humidity conditions. The present study tested the adequacy of the three models on the cockroach’s moist and dry cells by determining whether the specific predictions about the temperature-dependence of the humidity responses are indeed observed. While in previous studies stimulation consisted of rapid step-like humidity changes, here we changed humidity slowly and continuously up and down in a sinusoidal fashion. The low rates of change made it possible to measure instantaneous humidity values based on UV-absorption and to assign these values to the hygroreceptive sensillum. The moist cell fitted neither the mechanical hygrometer nor the evaporation detector model: the temperature dependence of its humidity responses could not be attributed to relative humidity or to saturation deficit, respectively. The psychrometer model, however, was verified by the close relationships of the moist cell’s response with the wet-bulb temperature and the dry cell’s response with the dry-bulb temperature. Thus, the hygroreceptors respond to evaporation and the resulting cooling due to the wetness or dryness of the air. The drier the ambient air (absolutely) and the higher the temperature, the greater the evaporative temperature depression and the power to desiccate.     

Low rates of change enhance effect of humidity on the activity of insect hygroreceptors

The inability to measure humidity during stimulation has so far prevented us from understanding the contribution of moist cells and dry cells to orientation in a gradient of humidity. The problem was solved in the present study by means of a UV-absorption hygrometer that made it possible to monitor humidity at a rate of 100 Hz. The antennal moist and dry cells of the cockroach were exposed to humidities alternatively falling or rising at low rates between -1% RH s(-1) and +1% RH s(-1) (relative humidity). Impulse frequency of both types of cells depended simultaneously on instantaneous humidity and its rate of change. High frequencies of the moist cells signal high humidity. But at a given humidity, the response frequency is higher still when humidity is also rising. Conversely, high frequencies of the dry cell signal low humidity, and frequency is higher still at a given humidity when humidity is also falling. These responses ensure that the cockroach spent a minimum time in environments where desiccation or hydration occur and may thus protect the animal from emerging accidentally from under cover into moving air. In the constant-humidity retreat of the cockroach, fluctuating or even drifting discharge frequencies could serve as an early warning: return!     

Increased turning per unit distance as an area-restricted search mechanism in a pause-travel predator, juvenile plaice, foraging for buried bivalves

During searching, discovery of a prey patch by juvenile plaice Pleuronectes platessa was associated with a change from extensive to intensive search behaviour several moves before an attack on a prey. Intensive search behaviour was characterized by reduced distance of moves, a greater rate of turning per unit distance and shorter pauses between moves. The increase in turn rate was associated with area-restricted seaching, while a decrease in distances moved suggests that plaice search more efficiently for prey when stationary than while moving. The klinokinetic mechanism that appears to regulate search behaviour in juvenile plaice should allow efficient exploitation of a range of prey distribution patterns based on localized cues alone. Such a mechanism is especially useful to a migratory predator, like plaice, whose foraging is subject to time constraints imposed by tidally available feeding areas.     

Gas exchange pattern in the two‐spot ladybird beetle,Adalia bipunctata,differs in dry vs. moist air

Gas exchange patterns in the ladybird beetle, Adalia bipunctata (L.) (Coleoptera: Coccinellidae), were investigated using an infrared gaseous analyser (IRGA) and a coulometric O₂ respirometer (manometric–volumetric system). Before testing, the beetles were kept either in dry (dehydrated) or moist (hydrated) conditions for 1 day. Their subsequent gas exchange patterns did not depend on their state of humidity but rather were controlled by the humidity of the insect chamber during gas exchange measurement. If this chamber contained dry air, the beetles exhibited CO₂ release by burst, which we interpreted as cyclic gas exchange (CGE) with inter‐burst periods, but if the chamber was switched to contain moist air, then cyclic CO₂ release was soon abandoned and a pattern of continuous gas exchange appeared. Measurements with the coulometric respirometer in moist air showed that continuous gas exchange was often associated with weak abdominal pulsations, which we interpreted as active ventilation. Their metabolic rate was lower during gas exchange cycles than during continuous gas exchange. We revealed that in the ladybird beetle metabolic rate increased in moist air when the gas exchange pattern transitioned from cyclic to continuous.     

Turning field size and its effects upon computer-simulated klinotactic orientation

The turning field is defined in the context of klinotaxis as the angular region(s) into which an organism may direct itself at any point in time and space while orienting within a stimulus gradient. The turning field size determines the size distribution of turns an organism can make during klinotaxis. Changes in turning field size affect the efficiency of klinotactic source location as measured by computer simulations of ideal behaviors. The optimal field size lies between 90 and 150 degrees. Turning field size also affects the appearance of search paths made by organisms locating an attractant source. The significance of turning field size is discussed and the described klinotactic model is proposed as a predictive model for orientation research.     

Temperature-dependent dry receptor on antenna ofPeriplaneta. Tonic response

Summary Identified as a dry receptive unit by transitory impulse frequencies up to about 200 imp/sec during rapid drops in humidity (Fig. 3) and which no other modality elicited, the dry unit on the antenna ofPeriplaneta americana is characterized by a regular (Fig. 4) and relatively high stationary impulse frequency (10–65 imp/s). Without exception the stationary discharge rate (tonic frequency) rose with falling values of stationary absolute humidity at constant temperature, and with rising values of stationary temperature of ambient air at constant humidity. Enthalpy and evaporation cooling appear to be ruled out as exclusive adequate explanations for this double dependence. No matter whether tonic frequency is plotted against absolute humidity based on either volume of moist air or weight of dry air, or against partial pressure of water vapor in ambient air, or the difference between saturation and partial vapor pressure, or against relative humidity, the dependence on the temperature is not eliminated (Figs. 5–7). Because a temperature range of about 20 °C and all humidities between 0 and 100% occupy the same large segment of the unit's tonic frequency spectrum (Fig. 8), the unit is termed bimodal.The author wishes to express indebtedness and gratitude to Prof. Dr. Hansjochem Autrum for backing the early stages of this project with the assistance of the Deutsche Forschungsgemeinschaft, to Prof. Dr. Helmut Altner for his prolonged support, to Dr. Hans-Jürgen Hinz for his helpful discussion of questions involving thermodynamics, to Prof. Dr. Friedrich Earth for helpful criticism of the text, and to Miss Christel Praeg for tireless technical assistance.     

Humidity behavior and the identification of hygroreceptors in the adult mealworm, Tenebrio molitor

The humidity behavior of undesiccated and desiccated adult Tenebrio molitor was investigated at 72 and 92%r.h. The behavior of the sexes was analyzed separately. The moist responses of desiccated beetles were correlated with the distribution of smooth-surfaced peg organs by removing various antennal segments. The dry responses of undesiccated beetles were associated with grooved and flat-tipped peg organs by removing various types of antennal sensilla with a laser. On the basis of their morphology and the results of the behavioral experiments, it has been concluded that the smooth-surfaced, grooved, and flat-tipped peg organs are likely to function as humidity receptors. A behavioral model has been proposed to explain the humidity responses of the beetle.     

Effects of green tree retention, prescribed burning and soil treatment on pine weevil (Hylobius abietis and Hylobius pinastri) damage to planted Scots pine seedlings

1 Feeding damage and mortality caused to planted Scots pine seedlings by the pine weevils Hylobius abietis and Hylobius pinastri were studied on burned and unburned sites with 0, 10 and 50 m³ per hectare levels of green tree retention from the second to the fourth summer after logging and burning of the sites. 2 The rate of severe feeding damage to pine seedlings caused by pine weevils was higher on burned clearcut sites than on unburned ones, whereas burning did not increase the feeding damage rate on sites with groups of retention trees. The damage rate in the fourth summer was approximately the same on burned and unburned sites. 3 Pine weevil feeding was the major cause of mortality of freshly planted pine seedlings on unburned sites. On burned sites, mortality was higher than the rate of severe feeding damage, particularly in the second summer after burning, possibly owing to fungal attack and abiotic factors. 4 At a retention tree level of 50 m³, feeding damage to the seedlings was lower than on clearcuts and at a 10 m³ retention tree level. Furthermore, on sites with 50 m³ of retention trees, scarification of the soil was found to decrease feeding damage more effectively than on clearcuts and 10 m³ sites. If the seedlings were situated in the centre of scarified patches, scarification alone was as effective as insecticide treatment on unscarified soil for decreasing feeding damage and mortality. 5 The results suggest that when burning is applied as silvicultural treatment after clear‐cuts, retention of trees is recommended to reduce the damages caused by pine weevils on pine seedlings.     

Effects of green tree retention, prescribed burning and soil treatment on pine weevil (Hylobius abietis and Hylobius pinastri) damage to planted Scots pine seedlings

1 Feeding damage and mortality caused to planted Scots pine seedlings by the pine weevils Hylobius abietis and Hylobius pinastri were studied on burned and unburned sites with 0, 10 and 50 m³ per hectare levels of green tree retention from the second to the fourth summer after logging and burning of the sites. 2 The rate of severe feeding damage to pine seedlings caused by pine weevils was higher on burned clearcut sites than on unburned ones, whereas burning did not increase the feeding damage rate on sites with groups of retention trees. The damage rate in the fourth summer was approximately the same on burned and unburned sites. 3 Pine weevil feeding was the major cause of mortality of freshly planted pine seedlings on unburned sites. On burned sites, mortality was higher than the rate of severe feeding damage, particularly in the second summer after burning, possibly owing to fungal attack and abiotic factors. 4 At a retention tree level of 50 m³, feeding damage to the seedlings was lower than on clearcuts and at a 10 m³ retention tree level. Furthermore, on sites with 50 m³ of retention trees, scarification of the soil was found to decrease feeding damage more effectively than on clearcuts and 10 m³ sites. If the seedlings were situated in the centre of scarified patches, scarification alone was as effective as insecticide treatment on unscarified soil for decreasing feeding damage and mortality. 5 The results suggest that when burning is applied as silvicultural treatment after clear‐cuts, retention of trees is recommended to reduce the damages caused by pine weevils on pine seedlings.     

Do the level of energy reserves, hydration status and Borrelia infection influence walking by Ixodes ricinus (Acari: Ixodidae) ticks?

Ixodes ricinus horizontal movement within a humidity gradient and the influence of infection by Borrelia burgdorferi sensu lato (s.l.) on tick walking were investigated. Nymphs were placed within an arena containing a humidity gradient ranging from 45 to 95% relative humidity (RH). After 1 h of acclimation at 70% RH ticks were released so that they could either stay, or walk towards either the wet or the dry end. Their position was recorded 2 h post-release. Fat content was quantified and Borrelia infection was detected using real-time PCR and PCR followed by Reverse Line Blotting. Among the 1500 ticks tested, 29·85% were infected. More low-fat nymphs walked inside the arena than high-fat individuals. When nymphs walked, more low-fat ticks walked towards wetter than drier air, whereas more high-fat individuals walked towards drier than wetter air. Among high-fat nymphs, a lower proportion of Borrelia-infected ticks walked inside the arena compared to uninfected individuals, as though spirochetes manipulated their arthropod vector to stay. However, Borrelia infection had no effect on walking direction towards the dry or the wet end. Hence, it appears that I. ricinus nymphs walk horizontally over short distances within a humidity gradient depending on both energy resources and Borrelia infection.     

EFFECTS OF TEMPERATURE AND HUMIDITY ON THE CLOTHES MOTH LARVA, TINEOLA BISELLIELLA HUM. (LEPIDOPTERA)

Methods for breeding clothes moths with very little attention are described.
Experiments were made in which fasting clothes moth larvae were exposed to a number of temperatures (35, 30, 22 and 10° C.), and four humidities at each temperature (0, 30, 60 and 90 per cent.). The rate of loss of weight of the larvae was determined, and al 50 the amount of solid matter lost.
In moderately dry or moderately moist air, the larvae maintained their water balance, but in very dry air the proportion of solid matter in their bodies rase, and in moist air it fell. A considerable amount of dry matter was lost by excretion and the production of silk, in addition to metabolism. These processes were not affected by humidity.
The results suggest that the rate at which water is evaporated from the !arvae is proportional to the saturation deficiency of the air.     

Effect of inspired air conditions on exercise-induced bronchoconstriction and urinary CC16 levels in athletes

Injury to the airway epithelium has been proposed as a key susceptibility factor for exercise-induced bronchoconstriction (EIB). Our goals were to establish whether airway epithelial cell injury occurs during EIB in athletes and whether inhalation of warm humid air inhibits this injury. Twenty-one young male athletes (10 with a history of EIB) performed two 8-min exercise tests near maximal aerobic capacity in cold dry (4°C, 37% relative humidity) and warm humid (25°C, 94% relative humidity) air on separate days. Postexercise changes in urinary CC16 were used as a biomarker of airway epithelial cell perturbation and injury. Bronchoconstriction occurred in eight athletes in the cold dry environment and was completely blocked by inhalation of warm humid air [maximal fall in forced expiratory volume in 1 s = 18.1 ± 2.1% (SD) in cold dry air and 1.7 ± 0.8% in warm humid air, P < 0.01]. Exercise caused an increase in urinary excretion of CC16 in all subjects (P < 0.001), but this rise in CC16 was blunted following inhalation of warm humid air [median CC16 increase pre- to postchallenge = 1.91 and 0.35 ng/μmol in cold dry and warm humid air, respectively, in athletes with EIB (P = 0.017) and 1.68 and 0.48 ng/μmol in cold dry and warm humid air, respectively, in athletes without EIB (P = 0.002)]. The results indicate that exercise hyperpnea transiently disrupts the airway epithelium of all athletes (not only in those with EIB) and that inhalation of warm moist air limits airway epithelial cell perturbation and injury.     

Turning behaviour related to optical stimuli in Tenebrio molitor

The walking behaviour of the beetle Tenebrio molitor was studied on a compensating running globe under optical stimuli. The animals show a clear frequency distribution for the duration of continuous walking between abrupt stops. The turning behaviour after a light is switched on or off is correlated with these frequency distributions. The circadian rhythmic changes of activity and of the sensitivity of the eyes influence the turning behaviour. A model describes the phenomenon of ‘sensory storage’ as the result of an integration over the reaction probabilities given by the frequency distributions.     

Stomatal responses to changes in humidity in plants growing in the desert

Summary The stomata of plants growing in the Negev Desert, namely the stomata of the mesomorphic leaves of Prunus armeniaca, the xeromorphic stems of Hammada scoparia, and the succulent leaves of Zygophyllum dumosum, respond to changes in air humidity. Under dry air conditions diffusion resistance increases. Under moist air conditions diffusion resistance decreases. When the stomata close at low air humidity the water content of the apricot leaves increases. The stomata open at high air humidity in spite of a decrease in leaf water content. This excludes a reaction via the water potential in the leaf tissue and proves that the stomatal aperture has a direct response to the evaporative conditions in the atmosphere. In all species the response to air humidity is maintained over a period of many hours also when the soil is considerably dry. The response is higher in plants with poor water supply then in well watered plants. Thus for field conditions and for morphologically different types of photosynthesizing organs the results confirm former experiments carried out with isolated epidermal strips.     

Testing associational resistance against pine weevils mediated by Lasius ants attending conifer seedlings

Natural enemies attracted to plants may provide those plants with protection against herbivores but may also protect neighbouring plants, that is through associational resistance. Ant attendance may be an important mechanism for the occurrence of such effects because ants can reduce the damage caused by a wide variety of herbivorous insects. Ants have been shown, in a previous field experiment, to decrease the damage caused by the pine weevil, Hylobius abietis (L.) (Coleoptera: Curculionidae), a pest species that causes high seedling mortality in forest regeneration areas. In this study, we specifically tested whether seedlings planted close to ant‐attended seedlings experience associational resistance. We did this under laboratory conditions using the ant species Lasius niger (L.) (Hymenoptera: Formicidae). The feeding damage by pine weevils was significantly reduced on seedlings attended by ants. The neighbouring seedlings, however, did not experience associational resistance. Nevertheless, some associational effects were observed as the number of weevils recorded on both ant‐attended and neighbouring seedlings was significantly lower compared with ant‐excluded seedlings.     

Effect of Timber Condition on Parasitization of Pine Weevil ( Hylobius abietis L.) Larvae by Entomopathogenic Nematodes under Laboratory Conditions

The large pine weevil ( Hylobius abietis L.) is one of the most important pests in coniferous reforestation in Europe. Larvae develop in the stumps of recently felled trees; the emerging adults feed on the bark of seedlings and may kill them. The ability of the entomopathogenic nematodes Heterorhabditis megidis and Steinernema carpocapsae to invade pine weevil larvae in Sitka spruce ( Picea sitchensis ) buried in moist sand was evaluated. Overall, four times as many H. megidis as S. carpocapsae invaded pine weevil larvae. The two species of nematode differed in their response to timber condition. The number of S. carpocapsae invading pine weevil larvae was twice as high in billets inoculated with the wood-rotting fungus Phlebiopsis gigantea as in fresh timber, while the number of H. megidis invading was reduced by 25%. Invasion into non-feeding insects (larvae of the wax moth Galleria mellonella ) contained in timber disks was also affected by timber quality, indicating that nematode behaviour was affected directly by the physical or chemical condition of the timber, though trophically mediated effects may also have been involved.     

Alteration of Components of Leaf Water Potential and Water Content in Velvetleaf under the Effects of Long-Term Humidity Difference

Velvetleaf (Abutilon theophrasti Medik.) was grown in growth chambers set at 45 or 85% relative humidity at 30°C, CO₂ 350 microliters per liter and 1000 micromoles per square meter per second of photosynthetically active radiation. Soil water potential was maintained at −0.05 megapascal by subirrigation with half strength Hoagland solution. The third, fourth, and fifth leaves from the base of 21- and 25-day-old plants were used for pressure-volume measurements. Components of leaf water status including water potential (osmotic and potential associated with the apoplast), leaf water content (apoplasmic and symplasmic water), and elastic modulus of leaf tissue were determined. Results indicate: (a) persistent dry air generated leaves with lower water potential at a given relative water content than did humid air; (b) the higher total leaf water content in plants grown in dry air was related to an increase in apoplasmic water, whereas symplasmic water remained similar in both humidity treatments; (c) difference in leaf water potential between low and high humidity treatments was related to decreased potential associated with the apoplast but not to a change in cell wall elasticity.