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(A) View of a composed leaf of Utricularia inflata with eight visible traps. (B) A single trap of Utricularia inflata. The trigger hairs are situated on the door but are not visible on this picture.

(A) View of a composed leaf of Utricularia inflata with eight visible traps. (B) A single trap of Utricularia inflata. The trigger hairs are situated on the door but are not visible on this picture.

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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 en...

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... buckling scenario The body of an Utricularia trap ( Fig. 1) is a millimeter sized bladder made of two layers of cells, and is closed by a door articu- lated around hinges. Specific glands pump water out of the trap, decreasing its volume. 4,5 Due to the geometry of the trap body, this transformation is smooth and continuous, 2,6 and entails a progressive lowering of the inner pressure. Since ...

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... The traps have the potential to capture and preyed on multiple animals one after the other and multiple prey animals can be captured within a single suction swirl [33]. In Utricularia, no morphological change or growth is required before a second capture [34]. These properties advocate the larvivorous potential of genus Utricularia which would be used as a biological control agent for medically important mosquito larvae. ...
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Background The carnivorous genus Utricularia also includes aquatic species that have the potential to trap a wide range of prey, leading its death due to anoxia. However, the effectiveness of such an approach with carnivorous plants for vector control has not been evaluated in Sri Lanka. Methods Early instar (i & ii) and late instar (iii & iv) larvae of Aedes aegypti were exposed to locally found bladderwort ( U. aurea Lour and Utricularia sp.). The experimental design was set with 10 larvae (both early and late instars separately) in 250 mL of water with bladderworts containing approximately 100 bladders in plant segments of both species, separately. Each treatment and control were repeated 50 times. The survival status of larvae was recorded daily until death or adult emergence. The larvae found whole or partially inside the bladders were attributed to direct predation. The Cox-regression model and Mantel-Cox log rank test were carried out to assess the survival probabilities of larvae in the presence of two bladderworts separately. Results The highest predation was observed when using early instar larvae in both U. aurea (97.8%) and Utricularia sp. (83.8%). The mortality caused due to predation by U. aurea was observed to be significantly higher according to the Mantel-Cox log-rank test (HR = 60.71, CI; 5.69–999.25, P = 0.004). The mortality rates of late instar stages of Ae. aegypti were observed to be lower in both U. aurea (82.6%) and Utricularia sp. (74.8%). Overall, the highest predation efficacy was detected from U. aurea (HR = 45.02; CI: 5.96–850.51, P = 0.017) even in late instar stages. The results suggested the cumulative predation in both plants on Ae. aegypti larvae was > 72%. Conclusions Utricularia aurea is a competent predator of Ae. aegypti larvae. Further, it is recommended to evaluate the feasibility of this plant to be used in the field as a control intervention in integrated vector management programmes.
... Among carnivorous plants, the largest (around 220 species) and one of the most cosmopolitan genera is Utricularia, which includes terrestrial, epiphytic and aquatic species [12]. The traps, which are characteristic of the genus, are complex structures usually 1 to 6 mm long [13] that employ a complex suction mechanism to capture and digest small invertebrates [14]. Inside the bladder traps, there are two type of glands. ...
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Background: The genus Utricularia belongs to Lentibulariaceae, the largest family of carnivorous plants, which includes terrestrial, epiphytic and aquatic species. The development of specialized structures that evolved for carnivory is a feature of this genus that has been of great interest to biologists since Darwin's early studies. Utricularia gibba is itself an aquatic plant with sophisticated bladder traps having one of the most complex suction mechanisms for trapping prey. However, the molecular characterization of the mechanisms that regulate trap development and the biophysical processes involved in prey trapping are still largely unknown due to the lack of a simple and reproducible gene transfer system. Results: Here, we report the establishment of a simple, fast and reproducible protocol for genetic transformation of U. gibba based on the T-DNA of Agrobacterium tumefaciens. An in vitro selection system using Phosphinotricin as a selective agent was established for U. gibba. Plant transformation was confirmed by histochemical GUS assays and PCR and qRT-PCR analyses. We report on the expression pattern of the 35S promoter and of the promoter of a trap-specific ribonuclease gene in transgenic U. gibba plants. Conclusions: The genetic transformation protocol reported here is an effective method for studying developmental biology and functional genomics of this genus of carnivorous plants and advances the utility of U. gibba as a model system to study developmental processes involved in trap formation.
... Bladderworts (Utricularia spp., Lentibulariaceae, Lamiales) have suction traps ("bladders") for catching invertebrates (i.e., Mette et al., 2000;Alkhalaf et al., 2009;Płachno et al., 2014Płachno et al., , 2015Darnowski et al., 2018;Płachno and Muravnik, 2018), and sometimes, also for bacteria and protozoa cultures (Sirová et al., 2009(Sirová et al., , 2018Płachno et al., 2012). They are considered to be the fastest predators in the plant kingdom (Vincent and Marmottant, 2011;Adamec, 2011aAdamec, ,b, 2012Vincent et al., 2011a,b;Poppinga et al., 2013Poppinga et al., , 2016Poppinga et al., , 2017. The Utricularia trap is a small hollow bladder with a trap entrance ( Figure 1A). ...
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Bladderworts (Utricularia, Lentibulariaceae, Lamiales) are carnivorous plants that form small suction traps (bladders) for catching invertebrates. The velum is a cuticle structure that is produced by specialized trichomes of the threshold pavement epithelium. It is believed that the velum together with the mucilage seals the free edge of the trap door and that it is necessary for correct functioning of the trap. However, recently, some authors have questioned the occurrence of a velum in the traps of the Utricularia from the various sections. The main aim of this study was to confirm whether velum occurs in the traps of the Utricularia species from the subgenera Polypompholyx, Bivalvaria, and Utricularia. The 15 species were examined from subg. Polypompholyx, subg. Bivalvaria, and subg. Utricularia. A velum was found in all examined Utricularia species. In the traps of the members of section Pleiochasia, there was an outer velum (forming a complete ring) and an inner velum. In the traps of Utricularia uniflora (Lasiocaules), there was only an inner velum. In these species, the formation of the velum was accompanied by intensive mucilage production, and as a result, when door was closed (set position), the mucilage and the velum touched the surface of the door. In members of both sections of Pleiochasia and Lasiocaules, the pavement epithelium had a more complicated structure (four to five zones) than in the members of the subgenera Bivalvaria and Utricularia in which three distinct zones occurred (an outer with a velum, a middle and an internal with the mucilage trichomes). Even in U. purpurea, where the threshold was a reduced pavement epithelium, it consisted of three functional zones and the presence of a velum. Two main types of velum have been proposed. A velum was present in Utricularia traps regardless of the trap type or the habitat (aquatic, epiphytic, and terrestrial species). We proposed broad definition of velum as cuticle membranes covered by mucilage; from a functional point of view, this definition is more useful and more reflects complexity of this structure.
... When trigger (sensory) hairs situated externally on the trap door are touched by prey the door opens, the animal is aspirated into the trap lumen and the door closes the trap watertight again, all completed within 3-5 ms. The complex door motion comprises a reversible buckling/unbuckling process associated with a convex/concave door curvature inversion (Joeyux et al., 2011;Singh et al., 2011;Vincent and Marmottant, 2011;Vincent et al., 2011a,b). The negative pressure is partly restored by pumping out of ca. ...
... It has recently been found that both cut off and intact traps can also fire spontaneously in the course of time, i.e., without any mechanical stimulation by prey (Adamec, 2011a,b;Vincent and Marmottant, 2011;Vincent et al., 2011a,b;Płachno et al., 2015). There was no quantitative difference between spontaneous and mechanically stimulated firings and subsequent resetting rates (Adamec 2011a,b;Płachno et al., 2015). ...
Article
Firing and resetting of aquatic Utricularia traps are associated with water flows and pressure changes. A negative pressure of ca. −0.16 bar is formed in reset traps, but its direct measurement is very difficult. We present a method of a gradual external application of negative pressure of −0.56 to −0.84 mbar s−1 through a fine capillary to cut off aquatic Utricularia traps to determine the critical negative pressure (CNP) at which the traps (located in air) fire and aspirate an air bubble. Using an electronic pressure sensor, we simulated the physiologically formed negative pressure needed for spontaneous trap firing in 15 aquatic Utricularia species of four generic sections. Mean CNP values ranged from −0.069 bar in giant traps of U. reflexa to −0.346 bar in U. dichotoma. The average in all 20 species or variants tested was −0.195 ± 0.018 bar, while that in 13 species or variants of the generic section Utricularia was −0.165 ± 0.015 bar and significantly differed from that of three populations of two species (U. dichotoma, U. volubilis) of the generic section Pleiochasia (-0.335 ± 0.006 bar). CNP differed significantly between giant and smaller traps of U. reflexa and young and old traps of U. vulgaris. Pooled data for 20 species or variants showed a significant negative linear correlation between trap length and CNP value. Within each species, high variability of the CNP was found: the lowest values were usually 2-3 times lower than the highest ones. This variability can represent three types of spontaneous firings described in the literature.
... At least 724 species of vascular plants use prey capture to improve their mineral nutrition; with about 228 species, Utricularia (bladderworts, Lentibulariaceae) forms the largest and most widely spread genus of carnivorous plants (McPherson, 2010). In Utricularia, prey is captured by means of sophisticated suction traps (Lloyd, 1942;Sasago and Sibaoka, 1985a, b;Adamec, 2011a;Vincent and Marmottant, 2011;Vincent et al., 2011a, b). ...
Article
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Some carnivorous plants trap not only small animals but also algae and pollen grains. However, it remains unclear if these trapped particles are useless bycatch or whether they provide nutrients for the plant. The present study examines this question in Utricularia, which forms the largest and most widely spread genus of carnivorous plants, and which captures prey by means of sophisticated suction traps. Utricularia plants of three different species (U. australis, U. vulgaris and U. minor) were collected in eight different water bodies including peat bogs, lakes and artificial ponds in three regions of Austria. The prey spectrum of each population was analysed qualitatively and quantitatively, and correlated with data on growth and propagation, C/N ratio and δ(15)N. More than 50 % of the prey of the Utricularia populations investigated consisted of algae and pollen, and U. vulgaris in particular was found to capture large amounts of gymnosperm pollen. The capture of algae and pollen grains was strongly correlated with most growth parameters, including weight, length, budding and elongation of internodes. The C/N ratio, however, was less well correlated. Other prey, such as moss leaflets, fungal hyphae and mineral particles, were negatively correlated with most growth parameters. δ(15)N was positively correlated with prey capture, but in situations where algae were the main prey objects it was found that the standard formula for calculation of prey-derived N was no longer applicable. The mass capture of immotile particles confirms the ecological importance of autonomous firing of the traps. Although the C/N ratio was little influenced by algae, they clearly provide other nutrients, possibly including phosphorus and trace elements. By contrast, mosses, fungi and mineral particles appear to be useless bycatch. Correlations with chemical parameters indicate that Utricularia benefits from nutrient-rich waters by uptake of inorganic nutrients from the water, by the production of more traps per unit of shoot length, and by the capture of more prey particles per trap, as nutrient-rich waters harbour more prey organisms. © The Author 2014. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For Permissions, please email: journals.permissions@oup.com.
... When trigger hairs situated on the trap door are touched by a prey species the door opens, the prey is aspirated into the trap lumen and the door closes again; all in under 5 ms. This process is caused by the reversible buckling of the door (Vincent and Marmottant, 2011;Vincent et al., 2011a,b). It has recently been discovered that the traps can also fire spontaneously, i.e., without any mechanical stimulation (Adamec, 2011a,b;Vincent and Marmottant, 2011;Vincent et al., 2011a,b). ...
... This process is caused by the reversible buckling of the door (Vincent and Marmottant, 2011;Vincent et al., 2011a,b). It has recently been discovered that the traps can also fire spontaneously, i.e., without any mechanical stimulation (Adamec, 2011a,b;Vincent and Marmottant, 2011;Vincent et al., 2011a,b). Additionally, all recent studies indirectly support a purely physical (mechanical) rather than an electrophysiological mechanism of Utricularia trap triggering (Adamec, 2012). ...
... Both the formation of relatively large traps (up to 4-6 mm) in most aquatic Utricularia species and their ease of manipulation within their growing medium have meant that all studies on trap function have been confined to these species (Sydenham and Findlay, 1973;Sasago and Sibaoka, 1985;Adamec, 2011a,b;Adamec, 2011a,b;Singh et al., 2011;Vincent and Marmottant, 2011;Vincent et al., 2011a,b). Aquatic and/or amphibious Utricularia species represent only ca. ...
... When trigger hairs situated on the trap door are touched by a prey species the door opens, the prey is aspirated into the trap lumen and the door closes again; all in under 5 ms. This process is caused by the reversible buckling of the door (Vincent and Marmottant, 2011;Vincent et al., 2011a,b). It has recently been discovered that the traps can also fire spontaneously, i.e., without any mechanical stimulation (Adamec, 2011a,b;Vincent and Marmottant, 2011;Vincent et al., 2011a,b). ...
... This process is caused by the reversible buckling of the door (Vincent and Marmottant, 2011;Vincent et al., 2011a,b). It has recently been discovered that the traps can also fire spontaneously, i.e., without any mechanical stimulation (Adamec, 2011a,b;Vincent and Marmottant, 2011;Vincent et al., 2011a,b). Additionally, all recent studies indirectly support a purely physical (mechanical) rather than an electrophysiological mechanism of Utricularia trap triggering (Adamec, 2012). ...
... Both the formation of relatively large traps (up to 4-6 mm) in most aquatic Utricularia species and their ease of manipulation within their growing medium have meant that all studies on trap function have been confined to these species (Sydenham and Findlay, 1973;Sasago and Sibaoka, 1985;Adamec, 2011a,b;Adamec, 2011a,b;Singh et al., 2011;Vincent and Marmottant, 2011;Vincent et al., 2011a,b). Aquatic and/or amphibious Utricularia species represent only ca. ...
Article
Carnivorous plants typify a mixotrophic strategy where autotrophy is supported by predation on animals, achieved through fascinating morpho-physiological adaptations and unique mutualisms. Exploring such symbiotic interactions is pivotal to understand how carnivorous plants feed upon wide ranges of resources, by relying on symbiont-mediated digestion. This concept finds its extreme realization in Utricularia (bladderwort), the largest genus of carnivorous plants, where the formation of a symbiotic microbiota within the traps drives its trophic resources further, including algae and complex organic matter. The present review synthesizes the key aspects of Utricularia ecology, from evolution to habitat selection, looking into the fascinating complexity of the structure and the microbiome of their traps. It highlights the gaps and guts in the understanding of bladderworts, including promising applications based on models of the Utricularia-microbiome system and the possible development of biomimetic devices. From the environmental drivers through the solutions developed by these plants, the review offers a comprehensive view of their unique ecology, which could be factually defined a “mixotrophic omnivory”.
Article
Despite their lack of a nervous system and muscles, plants are able to feel, regulate flow, and move. Such abilities are achieved through complex multi-scale couplings between biology, chemistry, and physics, making them difficult to decipher. A promising approach is to decompose plant responses in different blocks that can be modeled independently, and combined later on for a more holistic view. In this perspective, we examine the most recent strategies for designing plant-inspired soft devices that leverage poroelastic principles to sense, manipulate flow, and even generate motion. We will start at the organism scale, and study how plants can use poroelasticity to carry information in-lieu of a nervous system. Then, we will go down in size and look at how plants manage to passively regulate flow at the microscopic scale using valves with encoded geometric non-linearities. Lastly, we will see at an even smaller scale, at the nanoscopic scale, how fibers orientation in plants' tissues allow them to induce motion using water instead of muscles.
Article
The carnivorous plant bladderwort exemplifies the use of accumulated elastic energy to power motion: respiration-driven pumps slowly load the walls of its suction traps with elastic energy (∼1 h). During a feeding strike, this energy is released suddenly to accelerate water (∼1 ms). However, due to the traps' small size and concomitant low Reynolds number, a significant fraction of the stored energy may be dissipated as viscous friction. Such losses and the mechanical reversibility of Stokes flow are thought to degrade the feeding success of other suction feeders in this size range, such as larval fish. In contrast, triggered bladderwort traps are generally successful. By mapping the energy budget of a bladderwort feeding strike, we illustrate how this smallest of suction feeders can perform like an adult fish.