Water balance in Collembola and its relation to habitat selection: Water content,haemolymph osmotic pressure and transpiration during an instar

In the two collembolan species Orchesella cincta and Tomocerus minor the water content, haemolymph osmotic pressure and transpiration rate fluctuate with the feeding rhythm during each instar. The changes in water content, however, are due to changes in dry weight, because the absolute water weight stays constant during the instar.The intake of food is probably the cause of the increase in haemolymph osmotic pressure. Increase of osmotically active substances in the blood and/or blood volume reduction may be responsible for the rise in osmotic pressure. This change in osmotic pressure in turn may affect the responsiveness of the animals to water as well as their feeding behaviour.Changes in the epicuticle and in epidermal cell membranes may cause changes in the rate of transpiration. The high rate observed during ecdysis and during the mid-instar may explain the behaviour of the animals in varied water conditions.Dehydration during the instar causes an equivalent rise in osmotic pressure for both Tomocerus minor and Orchesella cincta. The water loss appears to involve the haemolymph. The physiological state of the animal does not influence the rise in osmotic pressure. There are no signs of any osmoregulation in the two species.     

Size-specific mortality patterns in two species of forest floor Collembola

Summary Forest floor populations of Orchesella cincta (L) and Tomocerus minor (Lubbock) (Collembola; Entomobryidae) have been sampled over two generations and from length frequency distributions survivorship curves were constructed for each generation. Size-specific mortality rates were computed from size-specific survivorship data. The results reveal an asymmetrically U-shaped mortality pattern, with disproportionally high juvenile mortality. The pattern changes from one generation to the next, in particular the post-hatching mortality. Evidence from behavioural ecological research on a Collembola-specialized ground beetle is cited in favour of the hypothesis that size-specific mortality patterns are moulded by size-specific predation.     

Aggregation pheromones in Collembola

The existence of aggregation pheromones is demonstrated in the Collembola Orchesella cincta, Orchesella villosa, and Tomocerus minor. Places conditioned by these animals can be considered as pheromone-sources, causing aggregation. Although for Tomocerus minor perception is not strictly species specific, all species investigated appear to be more responsive to their own pheromone than to those produced by other species. Both males and females appear to produce and perceive the aggregation pheromone. The distant action of the pheromone points to an olfactorially acting pheromone, in the perception of which the antennae play an important rôle. The behavioural reaction of the perceiving animals is orthokinetic.     

Interspecific and intraspecific selection by the predator Notiophilus biguttatus F. (Carabidae) concerning two collembolan prey species

Summary Observations on predation by the diurnally active predator Notiophilus biguttatus F., show that locomotory activity of the prey favours capture. Of the two prey species studied, Orchesella cincta and Tomocerus minor, the former is by far the most active one during the day. Since the moulting rhythm causes only part of a group of O. cincta to be active, predation towards this species is selective: active specimens are preferred. Tomocerus minor is almost inactive during the day (but may be activated by disturbance from the predator), which inhibits such a selection. Orchesella cincta also shows, when attacked, a greater escape ability than T. minor. So, inactive O. cincta are captured less easily than T. minor. Thus, locomotory activity and escape ability both cause a variable preference of the predator. When a sufficiently high number of active O. cincta specimens is present, this species will be preferred; when the number of active O. cincta specimens is low, e.g. as a consequence of predation, T. minor might be preferred.     

Pitfall-trapping as a method for studying surface dwelling collembola

Summary Pitfall-traps are used to study the phenology and activity phenomena of four surface dwelling Collembola e. g. Entomobrya nivalis (Linné), Lepidocyrtus cyaneus Tullberg, Orchesella cincta (Linné) and Tomocerus minor (Lubbock).It is concluded that Entomobrya is active during the day, whereas Orchesella and Tomocerus are active during the night. As to Lepidocyrtus no definite conclusion can be drawn.A digging-in effect is demonstrated for Tomocerus minor. A negative effect of the roofs appears during the day for Entomobrya nivalis and Lepidocyrtus cyaneus.Weather conditions exert an influence upon these surface dwelling Collembola.

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Zusammenfassung Die Fallenfangmethode wurde angewandt zum Studium der Phänologie und der Aktivität von vier Collembolen-Arten der Bodenoberfläche, nämlich Entomobrya nivalis (Linné), Lepidocyrtus cyaneus Tullberg, Orchesella cincta (Linné) und Tomocerus minor (Lubbock). Entomobrya erwies sich als tagaktive Art, Orchesella und Tomocerus sind nachtaktive Tiere. Für Lepidocyrtus waren die Resultate nicht eindeutig.Eine Reaktion auf das Eingraben der Fallen wurde festgestellt für Tomocerus minor.Fallendächer haben einen negativen Effekt auf den Fang von Entomobrya nivalis und Lepidocyrtus cyaneus während des Tages.Die Aktivität dieser Collembolen der Oberfläche wird durch Witterungsverhältnisse beeinflußt.

     

Physiological absorption of liquid water by Collembola: Absorption by the ventral tube at different salinities

The ability to absorb solutions has been examined in the coxal vesicles of the ventral tube in two sympatric surface dwelling Collembola (Tomocerus sp. and Orchesella villosa) from a beech forest. The net influx of distilled water and different sodium chloride solutions was measured, followed by examination of the effective surface of the vesicles which contacts the medium. The transport rate decreased with increasing salinity of the medium. Orchesella always showed higher absorption rates than Tomocerus, if the values were related to unit surface area. However, considering that the effective surface area of the vesicles of Tomocerus is larger, the total absorption rate by the ventral tube of Tomocerus exceeded that of Orchesella. Calculating the increase in the total water content. Orchesella compensated for its deficit faster than Tomocerus, because Orchesella is smaller in total weight and water content. In most cases the efficiency of the absorptive epithelium decreased during an absorption cycle; this also occurred before moulting. Some hours after moulting, the absorption rates increased to their former level. Comparing the rates of transpiration, absorption by the ventral-tube vesicles, and drinking, confirmed the dominant role of the ventral tube in the water balance of Collembola. It is an important factor in the strategy of adaptation from the hypogaic to the epigaic life.     

Effects of food deprivation and type of prey on predation by Notiophilus biguttatus F. (Carabidae) on springtails (Collembola)

Summary With diurnally active predators like Notiophilus biquttatus F. food deprivation is involved in predation in two ways: as a consequence of food shortage (i.e., low prey density), and as a consequence of the night period. The pattern of food intake after deprivation at night has been studied with respect to two prey species and differently deprived predators. They prey species represent a locomotory active one (Orchesella cincta) and a locomotory inactive one (Tomocerus minor). It appears that the rate of predation after deprivation shows a distinct pattern, initially high and then slowing to a more or less constant value. This pattern is influenced by food deprivation and type of prey. The beetles preying on O. cincta consumed more prey than those preying on T. minor. The more deprived predators compensated for deprivation by a higher daily predation when O. cincta was the prey, but not when T. minor was. Consequences of these findings are discussed with respect to diet composition and functional response.     

Formation and ecological significance of aggregations in collembola

Summary Spatial distribution in the two collembolan species Tomocerus minor and Orchesella cincta is greatly influenced by the distribution of the environmental factors water and food. T. minor is totally restricted in distribution to water-saturated places, where it forms spaced-out aggregations. O. cincta assembles in water-saturated places during ecdysis and subsequent reproduction. This leads to dense contact-aggregations. Dispersal follows during the feeding period, probably caused by food shortage, presence of other species and/or saturated conditions in the aggregation site. After feeding, orientation toward water-saturated places occurs by means of orthokinetic reactions and the aggregations are reestablished. The effect of this periodical aggregation for the population is discussed.     

Bioinformatics and protein expression analyses implicate LEA proteins in the drought response of Collembola

Humidity has a large impact on the distribution and abundance of terrestrial invertebrates, but the molecular mechanisms governing drought resistance are not fully understood. Some attention has been given to the role of the heat shock response as a component of desiccation tolerance, but recent focus has been on the chaperone-like LEA (late embryogenesis abundant) proteins in anhydrobiotic animals. This study investigates the expression of putative LEA proteins as well as the heat shock protein Hsp70 during drought stress in soil and surface dwelling species of Collembola (springtails). In silico analysis of four EST candidates from two species of Collembola showed the presence of a Group 3 LEA protein in Megaphorura arctica. In common with other Group 3 LEA proteins, the new sequence is predicted to be 100% natively unfolded, with a strong degree of lysine and alanine periodicity and with a negative average hydrophobicity of −1.273. The sequence clusters with members of the Group 3 LEA in plants. Furthermore, cross-species Western blotting showed drought-induced expression of putative LEA proteins in six species of Collembola. In the surface dwelling species, Orchesella cincta, degree of dehydration and length of exposure correlated with level of putative LEA protein. Hsp70 was also found to increase in individuals of O. cincta and Folsomia candida that had been exposed to drought conditions for 6 days. These results show the presence of a LEA protein-coding region in Collembola, but also indicate that several proteins are involved in response to dehydration stress, including Hsp70.     

A comparative study of winter survival in two temperate Collembola

ABSTRACT.

• 1 We studied winter survival of two temperate collembolan species, Orchesella cincta (L.) and Tomocerus minor (Lubbock), that overwinter in adult and juvenile life stages.
• 2 A significant seasonal variation in cold hardiness, measured as lethal temperature after 24 h exposure, was found for both species. There was no difference between juveniles and adults in cold hardiness. O.cincta was more cold tolerant than T.minor.
• 3 A group of high and a group of low values for lethal temperatures could be distinguished.
• 4 Frost periods induced gut evacuation in O.cincta, but not in T.minor.
• 5 During frost periods no extra winter mortality was found in T.minor. This species moved down the soil profile as a response to a frost period without snow cover.
• 6 The winter mortality of O.cincta was difficult to estimate, since part of the population remained above ground in trees and escaped from sampling. The smallest size classes, which are restricted to the litter, had a lower winter mortality than T.minor.

     

Heavy-metal adaptation in terrestrial invertebrates: A review of occurrence,genetics, physiology and ecological consequences

1. The occurrence of genetic adaptation to heavy metals in natural populations of terrestrial invertebrates is evaluated from literature data. Five criteria for adaptation evidence are applied, with concepts from ecotoxicology, ecology, life-history theory and quantitative genetics.2. There is strong evidence for the occurrence of adaptation in natural populations of the isopod Porcellio scaber (Isopoda), the springtails Isotoma notabilis, Onychiurus armatus and Orchesella cincta (Collembola), the blowfly Lucilia cuprina and the fruit fly Drosophila melanogaster (Diptera). Adaptation to metal-containing pesticides has been demonstrated in ticks (Acarina). Population divergence indicates acclimation or adaptation in many other species.3. Metal adaptation has been achieved within a few generations under laboratory conditions in some species; adapted populations occur at field sites that have been polluted for decades, or longer.4. Genetic variation for tolerance and life-history characteristics, allowing for adaptation, was quantified in a reference population of Orchesella cincta. Tolerance and life-history patterns in exposed field populations matched predictions from genetic variation.5. Adaptation involves modification and intensification of existing physiological mechanisms for metal assimilation, excretion, immobilization or compartmentalized storage. There are indications ofinter-population divergence in metal-binding proteins in a snail. In the fruit fly Drosophila melanogaster metal adaptation is achieved by duplication of the metallothionein gene.6. An altered life-history is often part of the complex adaptation syndrome. Metal-adapted invertebrates have a shorter life-cycle and a higher reproductive effort.7. Possible consequences of adaptation, consisting of costs of tolerance determined by genetic correlations, and probably of reduced genetic variation for tolerance and other features, are discussed. Reduced genetic variation is suggested by results for the springtail Orchesella cincta.8. The distinction between “costs of tolerance” on the one hand and linkage disequilibrium or direct selection for altered life-history patterns on the other hand is discussed.9. Species with high sensitivity (i.e. a low NOEC), that do not have populations maintaining sufficient genetic variation to evolve tolerance or modified life-history characteristics, or that have costly tolerance mechanisms, or both, are most at risk for extinction at sites with increasing metal pollution.10. Metal adaptation in terrestrial invertebrates appears to be of degree rather than of kind: indications for a specific metal-fauna, equivalent to metal-vegetation, are lacking.     

Relationship between saturation deficit and the survival and locomotory activity of surface dwelling Collembola

The springtails Tomocerus minor, Orchesella cincta and Isotoma viridis survive only in air with a saturation deficit near zero. Their survival time is inversely related to saturation deficit and not simply to the relative humidity. The different species show different reactions to the same saturation deficit values, which agree with the humidity preferences of the species in their natural habitats.Drought stimulates Collembola to start locomotory activity which may lead them to aggregate in optimal humidity conditions. Thus this reaction may play a part in increasing the survival rate of these animals.

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Zusammenfassung Die Beziehung zwischen Sättigungsdefizit (S.D.) und Überlebenszeit und zwischen S.D. und lokomotorischer Aktivität wurde an drei epedaphischen Collembolen-Arten untersucht. Die Überlebensdauer verhält sich umgekehrt proportional zum S.D., und steht in keinem direkten Verhältnis zur relativen Luftfeuchte. Die verschiedenen Arten weisen unterschiedliche Reaktionen zu den gleichen S.D.-Werten auf. Diese Reaktionen stimmen mit den Feuchte-Präferenzen der drei Arten in ihren natürlichen Habitats überein. Hohe S.D.-Werte führen zu lokomotorischer Aktivität, mittels welcher die Tiere optimale Feuchtigkeitsverhältnisse erreichen können.Diese Ergebnisse weisen auf einen ökologischen Überlebensmechanismus hin.

     

Microsatellite loci in the soil‐dwelling collembolan,Orchesella cincta

Three total genomic libraries of the springtail Orchesella cincta (Insecta; Collembola) were screened for the presence of microsatellites using phosphor and colour detection. PCR primers were successfully constructed for six dinucleotide, one trinucleotide, and one interrupted dinucleotide microsatellite repeat. None of the microsatellite arrays were longer than 11 repeat units. Individuals from three forests were investigated: five out of eight microsatellite markers were polymophic in all forests (2–5 alleles per locus) and the observed heterozygosity was 0.1–0.9 per locus.     

Water relations and photosynthetic water use efficiency as indicators of slow climate change effects on trees in a tropical mountain forest in South Ecuador

The effects of an increasing moisture on trees of the tropical species-rich mountain rain forest in the South Ecuadorian Andes was investigated, using the daily total water consumption (TWC) and the instantaneous water use efficiency (WUE, ratio of photosynthetic CO₂ uptake per water loss by transpiration) as ecophysiological indicators. Two canopy and one sub-canopy tree species, (Vismia tomentosa, Clusiaceae, an as of yet unknown Lauracee, and Spirotheca rosea, Bombacaceae) were the experimental objects. Seasonal changes as well as a long-term (18 months) trend of increasing precipitation caused an inverse reaction of the TWC of the trees. Because of a rather unlimited water supply to the trees from a permanently high water content of the soil, transpiration followed mainly the atmospheric demand of water vapor, and increasing moisture hence reduced water loss by transpiration. It was hypothesized that in spite of the reduction in transpiratory water loss photosynthetic carbon acquisition would be not or less affected due to an increase in water use efficiency. Concomitant measurements of photosynthetic net CO₂ uptake showed the expected increase of WUE in V. tomentosa and S. rosea, but no clear reaction of the Lauracee. Accompanying measurements of stem extension growth confirmed an undiminished growth of V. tomentosa and S. rosea but showed also suspended growth of the Lauracee during the wettest months. While TWC can be continuously monitored with the heat dissipation technique, WUE is determined by leaf porometry in campaigns for which access to the canopy is required. Simultaneous recordings of the gas exchange of leaves at 4 different positions in the crown of one of the experimental trees (V. tomentosa) showed the usability of the trait WUE in combination with the total daily water consumption as indicator set for assessing the response of trees to a subtly changing climate. However, not all tree species appear as likewise useful indicator trees.     

Transpiration as a function of soil temperature and soil water stress

An apparatus was developed for the measurement of transpiration rates of Trifolium repens. The transpiration rates were measured under controlled conditions of soil water stress and soil temperature. Other environmental parameters such as air temperature, relative humidity, light intensity and air speed were held constant. Diffusive resistances were calculated and stomatal aperture changes were recorded for all treatment combinations. A significant interaction between soil water stress and soil temperature was observed for stomatal closures. Stomatal closure was observed even in the so-called wet range of soil water stress. An increase in mesophyll resistance or incipient drying was observed for several treatment combinations. The mesophyll resistance was shown to increase as soil water stress increased.     

Studies on water transport through the sweet cherry fruit surface: IX. Comparing permeability in water uptake and transpiration

Water uptake and transpiration were studied through the surface of intact sweet cherry (Prunus avium L.) fruit, exocarp segments (ES) and cuticular membranes (CM) excised from the cheek of sweet cherry fruit and astomatous CM isolated from Schefflera arboricola (Hayata) Hayata, Citrus aurantium L., and Stephanotis floribunda Brongn. leaves or from Lycopersicon esculentum Mill. and Capsicum annuum L. var. annuum Fasciculatum Group fruit. ES and CM were mounted in diffusion cells. Water (deionized) uptake into intact sweet cherry fruit, through ES or CM interfacing water as a donor and a polyethyleneglycol (PEG 6000, osmotic pressure 2.83 MPa)-containing receiver was determined gravimetrically. Transpiration was quantified by monitoring weight loss of a PEG 6000-containing donor (2.83 MPa) against dry silica as a receiver. The permeability coefficients for osmotic water uptake and transpiration were calculated from the amount of water taken up or transpired per unit surface area and time, and the driving force for transport. Permeability during osmotic water uptake was markedly higher than during transpiration in intact sweet cherry fruit (40.2-fold), excised ES of sweet cherry fruit (12.5- to 53.7-fold) and isolated astomatous fruit and leaf CM of a range of species (on average 23.0-fold). Partitioning water transport into stomatal and cuticular components revealed that permeability of the sweet cherry fruit cuticle for water uptake was 11.9-fold higher and that of stomata 56.8-fold higher than the respective permeability during transpiration. Increasing water vapor activity in the receiver from 0 to 1 increased permeability during transpiration across isolated sweet cherry fruit CM about 2.1-fold. Permeability for vapor uptake from saturated water vapor into a PEG 6000 receiver solution was markedly lower than from liquid water, but of similar magnitude to the permeability during self-diffusion of ³H₂O in the absence of osmotica. The energy of activation for self-diffusion of water across ES or CM was higher than for osmotic water uptake and decreased with increasing stomatal density. The data indicate that viscous flow along an aqueous continuum across the sweet cherry fruit exocarp and across the astomatous CM of selected species accounted for the higher permeability during water uptake as compared to self-diffusion or transpiration.     

Diurnal Pattern of Transpiration,Water Uptake and Water Budget of Succulents with Different CO2 Fixation Pathways

The water fluxes and the CO₂ exchange of three leaf succulents, Othonna opima, Cotyledon orbiculata and Senecio medley-woodii, with different leaf anatomy, growth form and CO₂ fixation pathways (C₃, CAM) were monitored with a gas exchange cuvette which was combined with a potometric system to quantify water uptake. Measurements, which are primarily valid for plants with a sufficient water supply, were made during 6 to 10 consecutive days under constant experimental conditions. Water uptake for 24 h exceeded water loss by transpiration only for a S, medley-woodii plant with 10 expanding but only 7 mature leaves. In this case the gained water evidently is put into leaf expansion. All other plants showed balanced transpiration and water uptake rates. O. opima and C. orbiculata have a similar life form, similar water storage volumes and the same natural habitat but their diurnal water uptake patterns differ significantly. In the C₃ plant O. opima water uptake increased when the transpiration increased or transpiration rates were higher than uptake rates and vice versa. On the contrary the CAM plant C. orbiculata transpired during the dark period at constant or decreasing rates but showed steadily increasing uptake rates. Senecio medley-woodii- and C. orbiculata are CAM plants with similar diurnal water uptake patterns with its maximum in uptake during or towards the end of the CO₂ dark fixation period. Water uptake of C. orbiculata was at its minimum at the end of the light period despite transpiration being maximal. The results were discussed considering the different CO₂ fixation pathways. In the investigated CAM succulents, C. orbiculata and S. medley-woodii, the CAM influenced water uptake throughout the whole day and not only during the CO₂ dark fixation period.