Functional characteristics of traps of aquatic carnivorous Utricularia species

Firing and resetting of traps in aquatic Utricularia species are associated with water flow and trap volume changes. In this study, trap thickness was used as a measure of water flow and was monitored automatically using an electronic position sensor. The basic characteristics of mechanically stimulated firing and resetting were measured in isolated traps from 13 aquatic Utricularia species and in two trap size categories in Utricularia reflexa. This allowed to study the relationship between these trap functions and trap thickness and length (criteria of trap size). Additionally, the characteristics of spontaneous firings (without any mechanical stimulation) were compared for U. reflexa traps fed or denied prey during a 1-day period. On the absolute scale, the 13 Utricularia species differed considerably in their firing and resetting rates. Significant interspecific differences were also found in the magnitude of firing (in total 3.7–4.2 times) and resetting rates (10–24 times) per unit trap thickness or length. Overall, traps of Utricularia australis, Utricularia stellaris and Utricularia inflata showed the greatest firing and resetting rates. The relative magnitude of firing per unit trap thickness or length showed a highly significant negative correlation with both trap thickness and length and the same also held for the relative resetting rates. Smaller and narrower traps are thus relatively more effective at trap firing and resetting than larger traps. Neither firing nor resetting characteristics were significantly different between unfed and prey-fed traps of U. reflexa and this was also true for the occurrence of spontaneous firings. A strict linear resetting rate, without any lag-period, was found during the first 3 min after trap firing in U. reflexa. This suggests that water is pumped out of the trap continuously and probably recirculates. Given the concepts of spontaneous firing and water recirculation, an ecological model based on the literature data has been devised to quantify the daily N and P gain from the ambient water by Utricularia traps devoid of animal prey. The model shows that the total N and P content estimated in the trap fluid is too high to be accumulated from only the ambient water. This implies that the prey-free traps do not take up N or P from the trap fluid but rather exude a quantity of these nutrients to the fluid to support the microbial community. Therefore, the trap microorganisms behave more as parasites than commensals and represent an additional ecological cost for trap maintenance.     

Carnivorous Utricularia: The buckling scenario

We review recent results about the functioning of aquatic carnivorous traps from the genus Utricularia. The use of high speed cameras has helped to elucidate the mechanism at the origin of the ultra fast capture process of Utricularia, at a millisecond time scale. As water is pumped out of the trap, pressure decreases inside the trap and elastic energy is stored due to the change of shape of the trap body. This energy is suddenly released when the trap is fired: the trap door undergoes an elastical instability: buckling, which allows its fast and passive opening and closure. This mechanism is used by Utricularia both to catch preys touching its trigger hairs and to fire spontaneously at regular time intervals. The results leading to this interpretation are reviewed and discussed and suggestions for further work are briefly presented.     

Oxygen concentrations inside the traps of the carnivorous plants Utricularia and Genlisea (Lentibulariaceae)

BACKGROUND AND AIMS: Species of Utricularia and Genlisea (Lentibulariaceae) are carnivorous, capturing small prey in traps which are physiologically very active, with abundant quadrifid and bifid glands. Traps of Utricularia have walls composed of two cell layers, and are filled with water. Diverse communities of commensal microorganisms often live inside the traps. Genlisea forms long, hollow subterranean traps of foliar origin, growing in anoxic wet substrate. Knowledge of the O(2) concentrations inside Utricularia and Genlisea traps is vital for understanding their physiological functioning and conditions for the life of commensals. To test the hypothesis that prey are killed by anoxia inside the traps, and to measure respiration of traps, [O(2)] was measured in the fluid in mature traps of these species. METHODS: Oxygen concentration and electrical redox potential were measured using a small Clark-type oxygen sensor and a miniature platinum electrode, respectively, in the fluid of excised and intact traps of six aquatic Utricularia species and in Genlisea hispidula traps. KEY RESULTS: Steady-state [O(2)] in the traps of both genera always approached zero (median 0.0-4.7 microm). The [O(2)] decreased after electrodes were inserted into Utricularia traps at a rate which ranged from 0.09 to 1.23 mm h(-1) and was lower in traps of irradiated and intact shoots with higher [O(2)] in shoot tissues. Redox potential ranged from -24 to -105 mV in the traps, confirming the very small or zero [O(2)]. CONCLUSIONS: Very small or zero [O(2)], effectively anoxia, is demonstrated in Utricularia and Genlisea traps. This is probably below the critical [O(2)] for prey survival, and causes captured prey to die of suffocation. Internal trap glands and trap commensals are considered to be adapted to facultative anoxia interrupted by limited periods of higher [O(2)] after firings.     

The comparison of mechanically stimulated and spontaneous firings in traps of aquatic carnivorous Utricularia species

Firing and resetting of traps in aquatic Utricularia species are associated with water flows and trap volume changes. In this study, trap thickness was used as a measure of water flow and was monitored automatically using an electronic position sensor. Isolated traps from three aquatic Utricularia species were monitored over the course of 1-2 days to verify spontaneous firings (without any mechanical stimulation) and describe their basic characteristics. Isolated traps of three Utricularia species were initially fired by mechanical stimulation and allowed to naturally reset within a period of 24-48 h. Within this resting period, spontaneous firings were found in the traps of all species and in two trap age categories of U. vulgaris. The timing of spontaneous firings was found to be irregular. Spontaneous firings ranged between 0.29 and 2.4 during the 24-h resting period and the mean time between two spontaneous firings was highly variable within each species (319-891 min). There was no quantitative difference between spontaneous and mechanically stimulated firings of the traps. Spontaneous firings could explain how phytoplankton or detritus enters traps even when no prey species are present.     

Food or sex; pollinator–prey conflict in carnivorous plants

Carnivorous plants potentially trap their own pollinators and it has been argued that considerable spatial separation of flowers and traps has evolved to protect pollinators. We investigated flower-trap separation of Drosera and Utricularia . Short Drosera had a greater element of floral–trap separation than tall Drosera . Such a relationship is unexpected for plants whose peduncles were evolved to protect their pollinators. Utricularia can not trap pollinators but this genus still produces exceptionally long peduncles. We propose that flower-trap separation evolved because carnivorous plants are often short and need to project their flowers well above ground level to make them more attractive to pollinators.     

Ultra-fast underwater suction traps

Carnivorous aquatic Utricularia species catch small prey animals using millimetre-sized underwater suction traps, which have fascinated scientists since Darwin's early work on carnivorous plants. Suction takes place after mechanical triggering and is owing to a release of stored elastic energy in the trap body accompanied by a very fast opening and closing of a trapdoor, which otherwise closes the trap entrance watertight. The exceptional trapping speed--far above human visual perception--impeded profound investigations until now. Using high-speed video imaging and special microscopy techniques, we obtained fully time-resolved recordings of the door movement. We found that this unique trapping mechanism conducts suction in less than a millisecond and therefore ranks among the fastest plant movements known. Fluid acceleration reaches very high values, leaving little chance for prey animals to escape. We discovered that the door deformation is morphologically predetermined, and actually performs a buckling/unbuckling process, including a complete trapdoor curvature inversion. This process, which we predict using dynamical simulations and simple theoretical models, is highly reproducible: the traps are autonomously repetitive as they fire spontaneously after 5-20 h and reset actively to their ready-to-catch condition.     

The smallest but fastest: Ecophysiological characteristics of traps of aquatic carnivorous Utricularia

Aquatic Utricularia species usually grow in standing, nutrient-poor humic waters. They take up all necessary nutrients either directly from the water by rootless shoots or from animal prey by traps. The traps are hollow bladders, 1–6 mm long with elastic walls and have a mobile trap door. The inner part of the trap is densely lined with quadrifid and bifid glands and these are involved in the secretion of digestive enzymes, resorption of nutrients and pumping out the water. The traps capture small aquatic animals but they also host a community of microorganisms considered as commensals. How do these perfect traps function, kill and digest their prey? How do they provide ATP energy for their demanding physiological functions? What are the nature of the interactions between the traps and the mutualistic microorganisms living inside as commensals? In this mini review, all of these questions are considered from an ecophysiologist''s point of view, based on the most recent literature data and unpublished results. A new concept on the role of the commensal community for the plants is presented.Key words: aquatic carnivorous plants, bladderwort, bladders, firing, resetting, enzyme secretion, water pumping, microbial commensals     

Aging of Utricularia traps and variability of microorganisms associated with that microhabitat

Various authors have described algae in aquatic Utricularia traps as commensals, as stress factors or as prey. This study examined the diversity and abundance of organisms (prey, algae, protozoa and bacteria) in the traps of aquatic Utricularia reflexa in relation to prey occurrence and trap age. The number of organisms increased with the trap age. In both young and old traps, phytoplankton dominated of all organisms found. In young traps, Scenedesmus spp. and Characiopsis sp. were the most abundant algae, while Scenedesmus spp. and the palmelloidal form of Euglena spp. dominated in the old traps. Most of the algal species found stayed alive in the trap environment. The number of living algae and ciliates inside the traps increased with the increasing trap age, too. As the number of Paramecium bursaria inside traps consistently increased with the trap age and number of bacteria, which serve as a food for them, ciliates can be regarded as commensals, but not as prey for the plant. The predominant organisms in the traps were those that can be considered either commensals or intruders, exceeding captured macroscopic prey.     

Ecological Traits of the Algae‐Bearing Tetrahymena utriculariae (Ciliophora) from Traps of the Aquatic Carnivorous Plant Utricularia reflexa

Trap fluid of aquatic carnivorous plants of the genus Utricularia hosts specific microbiomes consisting of commensal pro‐ and eukaryotes of largely unknown ecology. We examined the characteristics and dynamics of bacteria and the three dominant eukaryotes, i.e. the algae‐bearing ciliate Tetrahymena utriculariae (Ciliophora), a green flagellate Euglena agilis (Euglenophyta), and the alga Scenedesmus alternans (Chlorophyta), associated with the traps of Utricularia reflexa. Our study focused on ecological traits and life strategies of the highly abundant ciliate whose biomass by far exceeds that of other eukaryotes and bacteria independent of the trap age. The ciliate was the only bacterivore in the traps, driving rapid turnover of bacterial standing stock. However, given the large size of the ciliate and the cell‐specific uptake rates of bacteria we estimated that bacterivory alone would likely be insufficient to support its apparent rapid growth in traps. We suggest that mixotrophy based on algal symbionts contributes significantly to the diet and survival strategy of the ciliate in the extreme (anaerobic, low pH) trap‐fluid environment. We propose a revised concept of major microbial interactions in the trap fluid where ciliate bacterivory plays a central role in regeneration of nutrients bound in rapidly growing bacterial biomass.     

Enzyme production in the traps of aquatic <Emphasis Type="Italic">Utricularia</Emphasis> species

We studied the influence of habitat and increased mineral phosphorus and nitrogen loading on the extracellular activity of five selected hydrolases and pH in the trap fluid of the aquatic carnivorous plants, Utricularia vulgaris, U. australis, and U. foliosa (Lentibulariaceae). Enzyme activities in the trap fluid were determined using fluorometry. Phosphatase exhibited the highest activities in the traps of the European species as well as field-grown tropical U. foliosa. Trap enzyme production appeared to be uninfluenced by elevated dissolved mineral N or P concentrations both in the trap and ambient environment and thus, it seems to be constitutive. Enzyme activity in the trap fluid was determined by species and environmental conditions and varied significantly among sites within a single species. Trap fluid pH was between 4.2–5.1 in U. vulgaris and U. australis but between 5.7–7.3 in U. foliosa and seems to be regulated by the traps.     

Quantifying responses of dung beetles to fire disturbance in tropical forests: the importance of trapping method and seasonality

Understanding how biodiversity responds to environmental changes is essential to provide the evidence-base that underpins conservation initiatives. The present study provides a standardized comparison between unbaited flight intercept traps (FIT) and baited pitfall traps (BPT) for sampling dung beetles. We examine the effectiveness of the two to assess fire disturbance effects and how trap performance is affected by seasonality. The study was carried out in a transitional forest between Cerrado (Brazilian Savanna) and Amazon Forest. Dung beetles were collected during one wet and one dry sampling season. The two methods sampled different portions of the local beetle assemblage. Both FIT and BPT were sensitive to fire disturbance during the wet season, but only BPT detected community differences during the dry season. Both traps showed similar correlation with environmental factors. Our results indicate that seasonality had a stronger effect than trap type, with BPT more effective and robust under low population numbers, and FIT more sensitive to fine scale heterogeneity patterns. This study shows the strengths and weaknesses of two commonly used methodologies for sampling dung beetles in tropical forests, as well as highlighting the importance of seasonality in shaping the results obtained by both sampling strategies.     

Rapid turnover of traps in Utricularia vulgaris L.

Summary Utricularia vulgaris growing at Wicken Fen, England, showed rapid turnover of the trapping apparatus. New groups of leaves, each bearing many traps, were produced at a rate of 1.4–2.8 groups per day from April to September, 1987. Old leaves decayed at a comparable rate, so that individual leaves survived for less than 50 days. In July, trapping efficiency of individual bladders was greatest between 1 and 6 days of age and then declined rapidly. Very few traps more than 19 days old could capture prey and most were lost from the leaves within 32 days. The lifespan of traps was closely related to their size and position on the leaves. Because of the rapidity of changes in trap condition with age, trap age is a vital consideration in any studies of the functional ecology of U. vulgaris     

Trap style influences wild pig behavior and trapping success

Despite the efforts of many natural resource professionals, wild pig (Sus scrofa) populations are expanding in many areas of the world. Although many creative techniques for controlling pig populations are being explored, trapping has been and still is the most commonly used method of population control for many public and private land managers. We conducted an observational study to examine the efficiency of 2 frequently used trap styles: a small, portable box-style trap and a larger, semi-permanent, corral-style trap. We used game cameras to examine patterns of trap entry by wild pigs around each style of trap, and we conducted a trapping session to compare trapping success between trap styles. Adult female and juvenile wild pigs entered both styles of trap more readily than did adult males, and adult males seemed particularly averse to entering box traps. Less than 10% of adult male visits to box traps resulted in entries, easily the least percentage of any class at any style of trap. Adult females entered corral traps approximately 2.2 times more often per visit than box traps and re-entered corral traps >2 times more frequently. Juveniles entered and re-entered both box and corral traps at similar rates. Overall (all-class) entry-per-visit rates at corral traps (0.71) were nearly double that of box traps (0.37). Subsequent trapping data supported these preliminary entry data; the capture rate for corral traps was >4 times that of box traps. Our data suggest that corral traps are temporally and economically superior to box traps with respect to efficiency; that is, corral traps effectively trap more pigs per trap night at a lower cost per pig than do box traps. © 2011 The Wildlife Society.     

Determination of an Optimal Pitfall Trap Size for Sampling Spiders in a Western Australian Jarrah Forest

Pitfall trapping is a sampling technique extensively used to sample surface foraging invertebrates for biological diversity studies and ecological monitoring. To date, very few invertebrate studies have considered what trap size is optimal for sampling spiders. This study presents preliminary findings from a single short sampling period on the role of trap size in sampling spiders in a Western Australian Jarrah forest. Four different trap diameters (4.3, 7.0, 11.1 and 17.4 cm) were examined (4 trap sizes × 15 replicates = 60 traps). Two-way ANOVAs revealed no significant interaction effects between trap size or the spatial positioning of transects within the study site along which the pitfall traps were arranged. Post-hoc tests revealed abundance, family richness and species richness increased with increasing trap sizes for traps 7.0 cm. No significant differences in these dependent variables occurred between 4.3 and 7.0 cm traps, or for species richness between 11.1 and 17.4 cm traps. Determination of an optimal trap size was undertaken by bootstrapping and calculating species accumulation curves for increasing numbers of traps used. Three different criteria were considered: equivalent number of traps (15), standardized sampling intensity (cumulative trap circumference, approximately 207 cm) and standardized cumulative handling time (approximately 1 hour 17 minutes). The largest trap size (17.4 cm) was most efficient in terms of number of traps and trap circumference. For the same number of traps, it caught 19 species whereas all other trap sizes caught ten species. At the standardized circumference, it caught seven species whereas all other trap sizes caught five. For handling time, however, the two largest trap sizes (17.4 and 11.1 cm) were optimal. Both caught nine species whereas all other traps caught 相似文献   

Bladder function in Utricularia purpurea (Lentibulariaceae): is carnivory important?

Utricularia purpurea is a rootless, free-floating, aquatic, carnivorous plant. I quantified biomass investment in U. purpurea traps and determined when traps begin to function and what they trap in natural habitats. In the Everglades of south Florida, plants invest an average of 26% of their biomass in bladders, although bladder number varies among sites and over time. Leaves begin trapping as they mature, and on leaves one whorl older than the most recently matured leaves, almost 100% of bladders have allochthonous material. Despite the substantial investment in their biomass, bladders capture few aquatic microinvertebrates. Almost all mature bladders, however, have living communities of algae, zooplankton, and associated debris. These results support the hypotheses that the important association in U. purpurea bladders is a mutualism rather than a predator-prey interaction and that the major benefit to the plants from bladders is derived from this community.     

Secretion trap tagging of secreted and membrane-spanning proteins using Arabidopsis gene traps

Secreted and membrane-spanning proteins play fundamental roles in plant development but pose challenges for genetic identification and characterization. We describe a "secretion trap" screen for gene trap insertions in genes encoding proteins routed through the secretory pathway. The gene trap transposon encodes a beta-glucuronidase reporter enzyme that is inhibited by N-linked glycosylation specific to the secretory pathway. Treatment of seedlings with tunicamycin inhibits glycosylation, resulting in increased activity of secreted beta-glucuronidase fusions that result from gene trap integration downstream of exons encoding signal peptides. In the 2,059 gene trap lines that we screened, 32 secretion trap expression patterns were identified in a wide variety of tissues including embryos, meristems, and the developing vasculature. Genes disrupted by the secretion traps encode putative extracellular signaling proteins, membrane transport proteins, and novel secreted proteins of unknown function missed by conventional mutagenesis and gene prediction. Secretion traps provide a unique reagent for gene expression studies and can guide the genetic combination of loss of function alleles in related genes.     

Is GC bias in the nuclear genome of the carnivorous plant Utricularia?driven by ROS-based mutation and biased gene conversion?

At less than 90 Mbp, the tiny nuclear genome of the carnivorous bladderwort plant Utricularia is an attractive model system for studying molecular evolutionary processes leading to genome miniaturization. Recently, we reported that expression of genes encoding DNA repair and reactive oxygen species (ROS) detoxification enzymes is highest in Utricularia traps, and we argued that ROS mutagenic action correlates with the high nucleotide substitution rates observed in the Utricularia plastid, mitochondrial, and nuclear genomes. Here, we extend our analysis of 100 nuclear genes from Utricularia and related asterid eudicots to examine nucleotide substitution biases and their potential correlation with ROS-induced DNA lesions. We discovered an unusual bias toward GC nucleotides, most prominently in transition substitutions at the third position of codons, which are presumably silent with respect to adaptation. Given the general tendency of biased gene conversion to drive GC bias, and of ROS to induce double strand breaks requiring recombinational repair, we propose that some of the unusual features of the bladderwort and its genome may be more reflective of these nonadaptive processes than of natural selection.     

Monitoring stored-product pests in food processing plants with pheromone trapping,contour mapping,and mark-recapture

Distribution and movement patterns of several species of stored-product pests in a food processing plant were investigated. The objectives of this study were to determine the temporal and spatial variation in abundance of stored-product pests using pheromone traps; assess the effectiveness of trap type, location, and number on monitoring insect populations; and to evaluate the nature of pheromone trap capture hot spots by measuring patterns of insect movement. We determined that the distributions of Trogoderma variabile Ballion, Lasioderina serricorne (F.), Tribolium castaneum (Herbst), and Plodia interpunctella (Hübner) within the facility were typically clumped and that foci of high trap captures, based on visual observation of contour maps, varied among species and over time. Trap type and location influenced the number of T. variabile captured: traps on the floor and along walls captured more individuals than hanging traps and traps next to support pillars. T. variabile was the predominant insect pest at this facility and from mark-recapture studies, we found that individual beetles moved across multiple floors in the facility and from 7 to 216 m though the warehouse.     

Feeding ecology of a carnivorous bladderwort (Utricularia uliginosa,Lentibulariaceae)

The terrestrial carnivorous bladderwort, Utricularia uliginosa Vahl. (Lentibulariaceae) was studied to determine the species assemblage present in traps of these plants in situ across four sites over 15 months. The immediate soil environment was also sampled to determine the fauna present, and to compare the fauna present in traps with the fauna in the environment. The soil fauna consisted of 10 taxon types, which occupied either pelagic, epibenthic or interstitial microhabitats. All were found in traps of U. uliginosa, with the main prey being interstitial taxa followed by epibenthic and occasionally pelagic taxa. Numbers of individuals of the two most abundant soil taxa (nematodes, Elaphoidella) varied independently across the four sites over the 15 months of the study, as did numbers of Elaphoidella in the traps of U. uliginosa. Numbers of nematodes in the traps of U. uliginosa showed significant differences among sites, but not differences among times. Comparison of the trap fauna with the soil fauna revealed differences in relative abundance between soil samples and trap samples for two of the three taxa examined. There was an under‐representation of nematodes in the traps relative to numbers in surrounding soil. There was an over‐representation of the copepod Elaphoidella in the traps of U. uliginosa relative to numbers in soil at some of the times of sampling. Acarina were equally abundant in soil and trap samples. The patterns observed for Elaphoidella and nematodes may be due to selectivity in trapping by U. uliginosa, and/or differences in digestibility of the prey. Elaphoidella individuals were found to be attracted to U. uliginosa in a behavioural experiment. This may contribute to the over‐representation of Elaphoidella in the traps of U. uliginosa in the field.     

Defining the fire trap: Extension of the persistence equilibrium model in mesic savannas

Mesic savannas are dominated by trees that are strong resprouters caught in a frequent fire trap. Persistence within this fire trap has been described by a resprout curve of SizeNext ~ f(Pre‐fire size), defined by the Michaelis‐Menten function. A key feature of this resprout curve is a stable persistence equilibrium that represents the size of individual plants upon which a population will converge over successive inter‐fire time steps under a given fire regime. Here, we contend that such a resprout curve does not adequately describe resprout tree dynamics in frequently burnt mesic savannas because it is constrained to an asymptote. We propose a new framework for modelling the resprout curve, which recognizes that local environmental stochasticity and growth patterns can interact to change the growth response function entirely, and thus more readily reflect the range of feasible resprout responses. Importantly, we define an unstable equilibrium representing the size above which individuals have escaped the fire trap and explore mechanisms that can shift an individual from persistence to escape. Through a case study from northern Australia, we confirm that our framework provides a simple yet practical approach to defining these critical aspects of savanna tree growth dynamics: persistence and escape.