ABSTRACT
SUCTION FEEDING IN THE CARNIVOROUS BLADDERWORT UTRICULARIA
Suction feeding is an important feeding mode in aquatic organisms and
is used across a considerable size range, from tadpoles to whales. Our current understanding is based on how adult fish feed and suggests that
suction feeding is not effective for organisms just a few millimeters in size. All suction feeders have to overcome the inertial and viscous forces exerted by the water when sucking in water plus prey, yet only the inertial forces contribute to prey capture, while viscous forces contribute just to the cost and reduce the effectiveness of prey capture. Large predators do not need to complete their suction strikes as quickly as small predators because the contribution of viscous forces is low. We therefore predicted that (1) small suction feeders complete feeding events more quickly than large suction feeders, and that (2) smaller suction feeders approach the lower size limit and hence cannot generate the same high flow speeds as larger suction feeders. We focused on two species of the aquatic carnivorous plant bladderwort, Utricularia gibba and U. vulgaris , that capture zooplankton in traps that are just 1-5 mm long. We quantified the movements of the bladders during feeding strikes and their peak flow speeds. We found that bladderwort feeding strikes are much briefer than those of adult fish, and that the smaller bladderwort species, U. gibba , generates slower flows than the larger U. vulgaris, suggesting that U. gibba feed near the lower size limit .
Matthew David Brown May 2016
SUCTION FEEDING IN THE CARNIVOROUS BLADDERWORT UTRICULARIA
by Matthew David Brown
A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science in Biology in the College of Science and Mathematics California State University, Fresno May 2016 APPROVED For the Department of Biology:
We, the undersigned, certify that the thesis of the following student meets the required standards of scholarship, format, and style of the university and the student's graduate degree program for the awarding of the master's degree.
Matthew David Brown Thesis Author
Ulrike K. Müller (Chair) Biology
Otto Berg Chemistry
John V. H. Constable Biology
For the University Graduate Committee:
Dean, Division of Graduate Studies AUTHORIZATION FOR REPRODUCTION OF MASTER’S THESIS
X I grant permission for the reproduction of this thesis in part or in its entirety without further authorization from me, on the condition that the person or agency requesting reproduction absorbs the cost and provides proper acknowledgment of authorship.
Permission to reproduce this thesis in part or in its entirety must be obtained from me.
Signature of thesis author: ACKNOWLEDGMENTS I would like to recognize the Biology Department and congratulate all
of their staff who make scientific research possible here at Fresno State. I am always taken aback by the amount of unbelievable talent and tireless effort
that make the graduate program both exciting and enlightening. My research on Suction Feeding in the Carnivorous Plant Utricularia is the product of not
only my hard work and determination but the unbelievable support available to me by the other professors and colleagues. First off I would like to acknowledge my graduate adviser Dr. Ulrike Muller and committee members, Dr. Otto Berg, and Dr. John Constable. I thank them for championing me as their graduate student and for helping me navigate through the process of becoming a scientist. Their scientific knowledge can only be surpassed by their support and encouragement and they were an integral part in my development as a scientist. I also appreciate Darius Khorshidchehr, David Ryan, Eshan Bhardwaj, Janneke Schwaner, Maxwell Hall, Nolan Avery, and Rayhan Kabir for their guidance and encouragement. Thank you Dr. Roi Holzman for your collaboration with us at the beginning of this project, for sharing your research experience on suction feeding, and for your expertise on PIV software. Thank you Dr. Steven Vogel for attending my first oral presentation at SICB; your presence made it a transcendent experience, and it will be something that I hold dear. I would also like to thank my parents; without them, none of this would be possible. To me this research represents not only a pursuit of knowledge toward understanding the world around us but also shows what can be achieved when great people collaborate and work together. TABLE OF CONTENTS Page
LIST OF TABLES ...... vi LIST OF FIGURES ...... vii
INTRODUCTION ...... 1 Suction Feeding – An Overview ...... 1
Suction Feeding Mechanics in Adult Fish ...... 2 Suction Feeding – Effects of Flow Regime ...... 3 Research Objectives, Aims and Hypothesis ...... 5 MATERIALS & METHODS ...... 7 Plant Husbandry ...... 7 Experimental Set-Up to Record Feeding Strikes ...... 9 Data Analysis ...... 13 RESULTS ...... 16 Plant Morphology ...... 16 Time Line of Suction Events ...... 17 Flow Generated During Suction Events ...... 19 DISCUSSION ...... 22
Comparison with Published Studies ...... 22 Main Conclusions and Future Directions ...... 24 REFERENCES ...... 25 APPENDICES ...... 28 APPENDIX A: INDEX OF KINEMATICS RECORDINGS ...... 29 APPENDIX B: INDEX OF FLOW RECORDINGS ...... 41
LIST OF TABLES
Page
Table 1 . Event during a suction event of Utricularia vulgaris versus Utricularia gibba ...... 19 Table 2 . Flow speeds and duration of the suction event determined by flow visualization for Utricularia vulgaris versus Utricularia gibba ...... 21
LIST OF FIGURES
Page
Figure 1 : Suction-feeding performance is determined by prey properties, predator traits, and their interactions (after Holzman et al., 2012)...... 3
Figure 2 : Photograph of a bladderwort strand ( Utricularia vulgaris )...... 4 Figure 3 : Left: Photograph of a bladderwort trap ( Utricularia gibba ). Right: schematic drawing of the trap...... 5
Figure 4 : Zooplankton being filtered from the CSU Fresno pond...... 9
Figure 5 : The Phantom V12.1 high-speed camera (right)...... 10
Figure 6 : Schematic top view of the macro photography stage (not to scale)...... 12 Figure 7 : Definition of gape (diameter of the mouth opening), funnel diameter (funnel only present in U. gibba), and bladder size (longest dimension of the trap)...... 13 Figure 8 : Spatial and time transects through the suction flow of a Utricularia gibba ...... 15
Figure 9 : Morphology of U. vulgaris (left) versus U. gibba (right)...... 17 Figure 10 : Time line of a suction event for U. gibba (left) and U. vulgaris (right). All time and flow speed values are averages...... 18 Figure 11 : Representative time transect of flow speed at half gape from the mouth for U. gibba (top) and U. vulgaris (bottom)...... 20
INTRODUCTION
Suction Feeding – An Overview Suction feeding is a common mechanism of prey capture used by many aquatic organisms and is the most common feeding mechanism in fish. The hydrodynamics of suction feeding have been studied extensively for decades. The focus of this research has been on adult fish, who have maximum gape sizes of 3 mm or greater (Holzman et al., 2008). There is much less experimental research on small suction feeders, such as larval fish, tadpoles, and bladderworts, who have gape sizes of 0.2 to 0.5 mm (Drost et al., 1988; Deban and Olsen, 2002; China and Holzman, 2014). Yet size is an important aspect of flow phenomena, including suction feeding, because flow phenomena strongly depend on size; in the case of suction feeding this is gape size. The relationship between fluid mechanics and size is formally described by the Reynolds number Re. Reynolds number is defined as the ratio of inertial to viscous forces and is calculated as flow speed multiplied by gape size divided by the kinematic viscosity of water. When Reynolds number is large (>1000), inertial forces dominate the flow. When Reynolds number is small (<100), viscous forces dominate the flow. Both inertia and viscosity play a significant role at intermediate Reynolds numbers (100
2004). During ram feeding, the fish’s entire body approaches the prey during the feeding strike (Wainwright et al., 2001). Some suction feeding specialists, in particular sea horses, pivot their heads to enhance their suction feeding (Roos et al., 2009). This mechanism seems to be effective even for newborn seahorses, whose gape is less than 0.5 mm (van Wassenbergh et al., 2009).
3
Figure 1 : Suction-feeding performance is determined by prey properties, predator traits, and their interactions (after Holzman et al., 2012).
Suction Feeding – Effects of Flow Regime What we know about large suction feeders does not directly apply to small suction feeders. We know that fish larvae and tadpoles use suction feeding, and that the smallest vertebrate suction feeders have gapes of less than 0.2 mm (Deban and Olsen, 2002). Recent studies have shown also that larval fish are not only ineffective suction feeders, but that their low capture success is due to the effects of flow regime, not lack of experience (China and Holzman, 2014). The smallest known suction feeders are carnivorous plants of the genus Utricularia , bladderwort (Figure 2). The bladderwort genus comprises more than 200 species (Müller and Borsch, 2005), many of whom are aquatic, capturing zooplankton in small underwater traps (Gordon and Pacheco, 2007). Bladderwort have garnered a lot of attention in the scientific 4 community because they have one of the smallest genome among angiosperms, a fact that might be explained by their particular prey capture mechanism (Ibarra-Laclette et al., 2013). Bladderwort traps are sealed by a trap door (Figure 3); they are set by osmotically pumping water out of the trap (Sagaso and Sibaoka, 1985), generating negative pressure inside the bladder (Singh et al., 2011). The traps are triggered when prey touches the trigger hairs at the trap door, which causes the trap door to collapse inward and the bladder to inflate, sucking in water and prey (Vincent et al., 2011; Singh et al., 2011). These capture events are extremely brief, less than 1 millisecond pass between the trap being triggered and the trap door opening (Vincent et al., 2011).
Figure 2: Photograph of a bladderwort strand ( Utricularia vulgaris ).
5
vestibule gape diameter trap door trigger hairs
funnel diameter
foot stalk
stolon
Figure 3: Left: Photograph of a bladderwort trap ( Utricularia gibba ). Right: schematic drawing of the trap.
Currently, there are no scientific explanations for why bladderwort generate such brief suction strikes. This master’s thesis is part of a larger project exploring a hydrodynamic explanation for the brevity of the suction event.
Research Objectives, Aims and Hypothesis In this project, we aim to better understand how small organisms use suction feeding to capture prey. Tadpoles (Deban and Olson, 2002), fish larvae (Drost et al., 1988) and bladderwort (Meyers, 1982) are known to use suction feeding. Yet the mechanics of suction feeding has only been explored in large predators, such as adult fish (Wainwright et al., 2007). Our current understanding implies that suction feeding is not an effective strategy for predators with gapes below 1 mm (Vogel, 1994). The first aim of this study is to study and identify the kinematics of a suction feeding event. We will identify the stages of a feeding strike, time sequences and duration of those stages, and which movements occur during 6 these stages (such as trap opening and closing). Bladderwort are among the smallest and fastest suction feeders (Vincent et al., 2011). They have underwater leaves that are modified into hollow bladders with a trap door to capture zooplankton (Müller et al., 2004). We will collect high-speed recordings to determine the sequence of events that constitute a suction feeding strike.
Our second aim is to describe the flow pattern of a suction feeding event generated by feeding strikes. We will quantify the flow speeds and flow event duration that occur during a feeding strike.
MATERIALS & METHODS
Plant Husbandry Two carnivorous plant species, Utricularia gibba and Utricularia vulgaris , were purchased from Carnivorous Plant Nursery (16128 Deer Lake Road Derwood, MD 20855; website: www.carnivorousplantnursery.com).
Handling and care instructions for carnivorous plants were obtained from the book “The Savage Garden” by Peter D’Amato (1998) and from Eshan Bhardwaj, a local expert on the care and cultivation of aquatic and terrestrial carnivorous plants. The plants were kept at the University greenhouse. The greenhouse provides some of the necessary external environmental controls, such as temperature and sunlight control. Shade is provided by placing cardboard boxes (60 X 40 cm) upright between the bladderworts and the greenhouse windows. Each box had 9 three-centimeter diameter holes evenly spaced at ten-centimeter intervals. This allowed attenuated sunlight to reach the bladderwort while suppressing growth of algae. Water acidity was monitored using a pH meter (API Pondcare); the optimal pH of for bladderworts is slightly acidic at pH 5.5 (D’Amato, 1998). When necessary, driftwood or additional sphagnum moss were added to the bladderwort aquarium to lower pH (D’Amato, 1998). Mineral content of water was monitored using a TDS meter “Total Dissolved Solids” electrical conductivity meter (HM Digital). Optimal mineral content is 100-180 ppm (D’Amato, 1998). The U. gibba specimen were cultivated in a large plastic cement- mixing tub (47 by 62 cm) with a layer of Long Fibered Sphagnum Moss 3 cm deep, in 8 cm of de-mineralized water. Between the sphagnum moss and the bladderwort was a plastic screen attached to a frame (51 X 39 cm). This 8 screen allowed the easy removal of the bladderworts from the aquarium whenever maintenance was needed. The U. vulgaris specimen were grown in
3.8-liter mason jars. The mason jars were filled 10 cm high with sphagnum moss and 13 cm of deionized water.
Carnivorous plants require live prey to ensure healthy growth (D’Amato, 1998). Zooplankton present in the pond on campus, outside the greenhouse, was used to feed our aquatic carnivorous plants. The zooplankton was harvested by inserting a 100 µm nylon mesh net to the output of the pond’s underwater pump. Within a period of 24 hours this setup filtered several thousand liters of pond water leaving a discharge of materials and small organisms in the nylon bag. The catch from the net were then lightly rinsed into a 19-liter bucket yielding a very dense supply of zooplankton. Two supplies of zooplankton were kept in stock, one with its prey concentration high and the other low. The container with the high prey concentration was used for experimentation, while the container with low prey concentration was used to cultivate more zooplankton, thus ensuring a sustained supply. The zooplankton cultures could be diluted over and over again into several aquaria. Adding a light bed (4 cm) of sphagnum moss and crumbled flakes of fresh-water fish food allowed the concentration of zooplankton to increase. Each aquarium was cleaned monthly as follows. Live zooplankton was captured in a 100 µm nylon mesh, the old water and sphagnum were discarded, and the contents of the net were rinsed back into a clean aquarium. A clean aquarium helps to ensure the health of the zooplankton, allowing them to maintain a high population in a limited space. All of the carnivorous plants and zooplankton were stored in the greenhouse. 9
Zooplankton was added to each aquarium containing Utricularia using disposable pipettes. The set-up to capture zooplankton is shown in Figure 4.
Figure 4 : Zooplankton being filtered from the CSU Fresno pond. Note: High Concentrations are needed in order to observe spontaneous feeding events with reasonable frequency.
Experimental Set-Up to Record Feeding Strikes
Bladder Preparation We excised bladders at the stolon using micro-surgery scissors (Roboz Surgical Instruments). We then used superglue to attach the stolen to a banjo wire (length of wire: 3 cm). We then placed the bladder in a glass cuvette (Starna 1.0 x 1.0 x 10.5 cm) by pushing the banjo wire end into a bed of Blu- Tack on the bottom of the glass cuvette.
Illumination for Recording High- Speed Video To record high-speed videos of feeding strikes, we illuminated the bladder using a high-powered red LED (1 Watt, 660 nm, with collimating optics). To record high-speed videos of the flow generated by feeding strike, we illuminated the bladder with a laser diode (Stocker-Yale/ Coherent Lasiris 10
TMFL) producing a thin sheet of uniform transverse intensity (200 mW at 810 nm).
High-Speed Filming A Phantom V12.1 high speed camera (Vision Research) was used to film all bladderwort suction events. The following frame rates were used: 10
000 fps (600 X 800 pixels), 18 000 fps (640 X 480 pixels), 28 000 fps (512 X 384 pixels), and 50 000 fps (320 X 280 pixels). The macro imaging system consists of a 24 mm objective lens (Nikon), mounted with a reversing ring on a 105 mm lens (Nikon). This combination yields a magnification of 105/24=4.4 with minimal distortion and great light-gathering power (~f/1.7). The filming setup is shown in Figure 5.
Figure 5 : The Phantom V12.1 high-speed camera (right). Note : The camera is mounted directly to a breadboard on a sled that allows translation parallel to the optic axis. The sample area (left is on a micrometer-driven mechanical stage able to move independently along the x, y, z axes. The leftmost stainless steel post mounts a 4-axis mechanical manipulator used to trigger the bladder, as observed from above with a stereo microscope. The rightmost stainless steel post holds the sheet-generating diode laser and optics. Power supplies toward the back of the photo are used to power the LED and laser. 11 Flow Visualization For Particle Image Velocimetry (PIV) and Particle Tracking Velocimetry
(PTV), a concentrated suspension of 6 µm nylon particles was added to the cuvette and illuminated with the laser. In order to minimize degradation of
the laser sheet and video image by light scattering, bladders were mounted in the corner closest to the laser and camera, but at least 2 gape diameters from
either cuvette wall. The body of the bladderwort is illuminated by scattered laser light. This strong and rapidly fluctuating background confounds automated velocimetry of the internal flows. Nevertheless, individual particles within the vestibule (Figure 3) can be tracked manually from frame to frame. For particle tracking, the bladder was sometimes illuminated from behind with a 0.5 W near-infrared LED (850 nm). Opaque particles (nylon, or metallic pigment) were introduced to the hood area by means of a pulled glass capillary micro-pipette, which also served as trigger probe. In this case the desired image plane was effectively selected by the optical depth of field, and by tracking only particles already within the bladder profile. The flow recording setup is shown in Figure 6.
Experimental Procedure to Trigger Capture Events All recordings were triggered manually after the suction event using the post-event trigger setting of the camera: the camera was set to record continually until the camera trigger is pushed. Once we pushed the camera trigger, the camera then stopped recordings and offered me to store all the images recorded up to the moment we pressed the camera trigger. We used two procedures to elicit suction events: artificial triggering and prey triggering. To trigger bladders artificially, we touched the trigger hairs at 12
Figure 6 : Schematic top view of the macro photography stage (not to scale). Note: Individual bladders are mounted by the stolon in a glass cuvette, which is then translated into the focus of a laser sheet oriented perpendicular to the plane of the illustration. The water is seeded with live prey and/or nylon particles. A microscope mounted above the cuvette is used to position the bladder and to steer a probe acting as artificial trigger. the trap door with a cat eye brow whisker. The whisker was mounted to a three-dimensional motion stage, which allowed me to move the whisker carefully and precisely. The whisker was oriented to minimally intrude into the laser light sheet during flow recordings. To trigger bladder with prey, we added ostracods to the water and recorded suction events using again the post-event trigger setting, which allowed me to save all the images recorded before we press the camera trigger. 13 Data Analysis
Morphology and Kinematics Video sequences were initially stored in a proprietary format of the camera. To process the recordings, all images were converted to a series of individual tiff files. From these images, we measured distances (in pixel units) with the measurement tools of open-source image manipulation software (GIMP 2.8) to calibrate our images. Then we calculated bladder dimensions and the timing of events from the known magnification and frame rate. Figure 7 shows how we determined bladder dimensions in Utricularia gibba .
bladder gape size
diamete funnel diameter
Figure 7 : Definition of gape (diameter of the mouth opening), funnel diameter (funnel only present in U. gibba), and bladder size (longest dimension of the trap).
From the high-speed recordings we determined the following time parameters: t 0 = time at which the trap door begins to move; t c = time at
which the door begins to close; t end = time at which the door is closed again. 14 Particle Image Velocimetry Each frame of a video sequence was converted to an uncompressed 12- bit tiff image. Velocity vectors were calculated using Open Source Image Velocimetry command-line tools compiled under Slackware Linux. Pixel intensities were first normalized by subtracting the lowest value from each image sequence (“subtract to minima” pre-processing). Then cross- correlations were obtained from 24 pixel x 24 pixel patches, repeated every 8 pixels in the x and y directions. Further processing of the resulting vectors was performed with Wolfram Mathematica: cross-correlations with a signal- to-noise ratio less than 1.05 were rejected; scale factors and symmetry axes were obtained from representative frames of the video and flow field; then speed distributions were computed as the sequence of vectors lying closest to the axial transect in space or half-gape point in time (Figure 8). In order to scale the spatial transects, each range value was divided by the measured gape diameter to yield dimensionless distance; the corresponding speeds were then scaled so as to set the measured value at 0.5 gape equal to unity. The choice of scaling factor was over-sensitive to random error if a single data point was used. Therefore, since the axial transects were well represented by a Gaussian curve, each was first fit to a Gaussian; this smoothed curve was then scaled so as to reach unity at 0.5 gape; then the scaling factor was applied to the measured speed values. From the flow field, we calculated time transects to track how flow speed changes over time at a point in space half a gape from the mouth (Figure 8). We used these time transects to estimate event duration. We defined event duration as the time it takes flow speed to reach half peak speed to the moment flow speed drops back down to half peak speed. 15
Figure 8: Spatial and time transects through the suction flow of a Utricularia gibba . Note: Yellow line: spatial transect along the central axis of the suction flow at the moment of maximum flow velocity. Yellow circle: position in the flow for the time transect at half a gape from the mouth in the center of the flow. Red arrows: flow velocity vectors as calculated by PIV. Visible in the image are the trap mouth and the cat whisker.
RESULTS
We recorded 51 suction events with U. vulgaris and 78 suction events with U. gibba . Of these recordings, 6 (21) recorded suction events were illuminated by LED to record kinematics and determine a timeline for U. vulgaris (U. gibba ); of these time-line recordings, 4 (2) were triggered artificially, 1 (18) were triggered with prey, and 1 (1) was spontaneous. We also made 45 (61) recordings of flow fields for U. vulgaris (U. gibba ); of these, 42 (55) were triggered artificially, and 3 (5) flow fields were triggered by prey, and none (1) were spontaneous.
Plant Morphology Our project focuses on two bladderwort species, Utricularia vulgaris and Utricularia gibba , who differ in their overall and in their bladder morphology. U. vulgaris has more bladders per strand, larger bladder, and a wider size range of bladders than U. gibba . Figure 9 shows the difference in bladder density per strand: U. vulgaris typically has twice to three times more bladders per strand than U. gibba . U. vulgaris bladders range in size from 0.7 to 2.5 mm, in contrast U. gibba bladders range in size from 0.5 to 1.5 mm. U. gibba has also a different bladder morphology from U. vulgaris . U. gibba has a prominent hood (vestibule) in front of the trap door; in contrast U. vulgaris has no such vestibule, putting its trap door close to the mouth. Both bladderwort species are aquatic. They not possess any root system, and instead float in the water. Both species grow continually, growing at one end and dying off at the other end.
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Figure 9 : Morphology of U. vulgaris (left) versus U. gibba (right). Photographs courtesy of Rayhan Kabir and Maxwell Hall.
Time Line of Suction Events The first aim of this study was to characterize the time line of the suction feeding event, in particular the stages of a feeding event, the time sequences and duration of those stages, and which movements occur during these stages. We found that a typical suction feeding event in the bladderwort U. vulgaris (U. gibba ) lasts 1.0±0.6 ms (n=21) (0.7±0.3 ms; n=6)
from the start of the trap door opening to the trap door being fully open and 2.3±1.3 ms (n=21) (1.9±0.7 ms; n=6) ms to the moment that the trap door is beginning to close (Figure 10). Prey is typically sucked into the bladder past the trap door between the time that the trap door is fully open and the trap door beginning to close. The suction event of U. vulgaris is longer than the one of U. gibba when comparing the time it takes the door to open completely (t-test: p=0.033). A typical time sequence for U. gibba is shown in Figure 10. 18
U. gibba Kinematic Timeline Initiation
Peak Flow Speed Duration Gape is 100% open
Gape closing Door begins to open Gape closed
-2 0 2 4 6 8 10 Fluid begins to move Prey past trap Flow Event door Duration
Prey begins to move Prey past gape
U. vulgaris Kinematic Timeline Initiation Peak Flow Speed Duration Gape is 100% open
Gape closing Door begins to open Gape closed
-2 -1 0 1 2 3 4 5 6 7 8 Fluid begins to … Prey begins to move Prey past gape Flow Event Duration Prey past trap door
Figure 10 : Time line of a suction event for U. gibba (left) and U. vulgaris (right). All time and flow speed values are averages.
19
Overall, we found significant differences between the two species only in the time it takes the door to fully open (t test, p = 0.02): U. vulgaris takes
longer to open its door than U. gibba (Table 1). Overall, the time to door closing completed is the most variable, as evident in the high standard
deviation for both species.
Table 1 . Event during a suction event of Utricularia vulgaris versus Utricularia gibba . event [ms] Utricularia Utricularia gibba vulgaris
Trap door begins to open 0 0 Trap door fully open 1.0±0.6 (38) 0.7±0.3 (42) Trap door begins to close 2.3±1.3 (36) 2.5±0.7 (28) Trap door closed 5.6±2.2 (36) 6.6±4.0 (23) Note: All data are given as mean±standard deviation (n number). The beginning of the door opening is defined as t 0=0 ms.
Flow Generated During Suction Events The second aim of this study was to describe the flow pattern of the suction feeding events. We found that U. gibba achieved lower peak flow speeds (determined at half gape from the mouth, as indicated in Figure 8)
than U. vulgaris : U. gibba achieved 0.15±0.04 m s-1 (n=23), U. vulgaris achieved 0.27±0.08 m s-1 (n=13) (Table 2). Furthermore, event duration (defined as the time from reaching half peak speed to dropping back to half peak speed) also differs between both species: U. gibba achieved 2.2±0.5 ms (n=23), U. vulgaris achieved 1.6±0.4 ms (n=11) (Table 2). In general, U. gibba achieved lower peak flow speeds and its suction events lasted longer. 20
The differences in peak speed and event duration foreshadowed the differences in flow velocity time course between the two species (Figure 11).
Below are two typical time courses of flow speed at half gape from the mouth for each species. U. gibba had a fast onset and slow offset. In contrast, U. vulgaris appeared to have a fast onset and offset. These observations based on flow speed (Table 2) are consistent with the kinematics observations
(Table 1). Flow speed and event duration are significantly different between the two species (t test, p>0.01).
Flow speed [m/s]
Flow speed [m/s]
Time [ms]
Figure 11 : Representative time transect of flow speed at half gape from the mouth for U. gibba (top) and U. vulgaris (bottom). Note: The 0 of the x axis is defined by the start of the recording, not the start of the suction event. 21
Table 2 . Flow speeds and duration of the suction event determined by flow visualization for Utricularia vulgaris versus Utricularia gibba . Parameter Utricularia vulgaris Utricularia gibba
Peak flow speed 0.27±0.08 m s-1 (n=13) 0.15±0.04 m s-1 (n=23)
Suction event duration 1.6±0.4 ms (n=11) 2.2±0.5 ms (n=23) Note: All data are given as mean±standrad deviation (n number). Suction event duration is defined as the time from reaching half peak flow speed to flow speed dropping back to half peak speed.
DISCUSSION
We found in our study that bladderwort can generate fast and brief suction flows. In both species examined in this study, Utricularia gibba and
U. vulgaris , the trap door opens fully within roughly 1 ms. Utricularia vulgaris opens its door more slowly, yet has a briefer suction event and reaches higher peak flow speeds. Event duration was defined as the time from reaching half peak flow speed to speed dropping back down to half peak speed. So having larger bladders enables U. vulgaris to generate briefer events that generate faster flows. These findings are consistent with our understanding of the hydrodynamics of suction feeding: if suction feeding is limited by viscous forces, than smaller suction feeders (traps with smaller gapes) should generate slower flows and longer feeding events (Drost et al., 1988).
Comparison with Published Studies
Suction Time Line – Event Duration Our findings concerning the suction event time line are largely consistent with existing literature. We found that Utricularia gibba took 0.7 ms to open its door, U. vulgaris took 1.0 ms. We found door opening times for two other species, U. inflata and U. stellaris . Vincent et al. (2011) found that the trap door of Utricularia inflata collapses within 1 ms. Singh et al. (2011) found that the trap door of Utricularia stellaris collapses within 0.5 to 0.7 ms. Vincent et al. (2011) find that the fast opening is due to the sudden buckling and collapse of the door – the door has a bistable shape and prey touching the 23
trigger hairs at the door causes the door to quickly switch from one to the other stable configuration.
Singh et al. (2011) also find that, while the opening of the trap door is not only fast but also very consistent in duration, the time to complete closure
is much more variable, ranging from 3 to 5.5 ms. This finding is consistent with our own data: we found that the average time from door opening to
closing is 6.6 ms for U. gibba (average of 23 values, range 3.3-21.0 ms) and 5.8 ms for U. vulgaris (average of 37 values, range 2.3-12.7 ms).
Suction Flow – Flow Speeds and Time to Peak Flow Our findings concerning the flow show lower flow values than previously reported. We found that Utricularia gibba reaches a peak flow speed of 0.15 m/s (average of 23 events; range 0.07-0.21 m/s) at half a gape from the mouth, U. vulgaris reaches 0.27 m/s (average of 13 events; range 0.13-0.37 m/s). Vincent et al. (2011) indicate that they recorded the velocity of tracer particles within half a gape of the mouth. In contrast, our values are the result of PIV analysis, not particle tracking and are consistently at half a gape from the mouth and are determined using particle image velocimetry, not particle tracking velocimetry. Given how quickly flow speed deteriorates with increasing distance from the mouth, it is not surprising that our values are lower (Roi Holzman, personal communication). We found that Utricularia gibba maintains a high suction flow for 2.2 ms (average of 23 events; range 1.4-3.1 ms), U. vulgaris does so for 1.6 ms (average of 11 events; range 0.7-2.1 ms). This event duration was defined as the time from reaching half of peak flow speed to the time that speed drops back to half peak speed, and hence should be shorter than the event duration 24
defined by the opening and closing of the trap door. In our experiments, the time from opening to closing the trap door was indeed roughly three times
longer than the event duration defined by flow speed. The only flow data available in the literature are form Vincent et al. (2011), suggesting that the
time from opening to closing the door in U. inflata is roughly 2 ms while the event duration defined by flow speed is less than 1 ms.
Main Conclusions and Future Directions Our study shows that bladderwort can create brief yet powerful suction flows to catch aquatic prey. Our findings are consistent with previously published studies on two different species ( Utricularia inflata and U. stellaris ) from the two species used in our study ( Utricularia gibba and U. vulgaris ). Our study documents for the first time the complete flow fields, providing rigorous data for a hydrodynamic analysis of the flow generated by bladderwort. In the future, the flow data from this study will serve to test hydrodynamic models about suction feeding. The data will help us to determine how small suction feeders can generate suction flows despite their small size. Hydrodynamic models predict that small suction feeders should not be able to generate fast suction flows, yet they clearly do. REFERENCES REFERENCES
China, V. and Holzman, R., 2014. Hydrodynamic starvation in first-feeding larval fishes. Proceedings of the National Academy of Sciences , 111(22), pp.8083-8088.
D’Amato, P., 1998. The savage garden. Cultivating Carnivorius Plants. Ten Speed Press, California.
Deban, S.M. and Olson, W.M., 2002. Biomechanics: suction feeding by a tiny predatory tadpole. Nature , 420(6911), pp.41-42.
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Holzman, R., Collar, D.C., Mehta, R.S. and Wainwright, P.C., 2012. An integrative modeling approach to elucidate suction-feeding performance. The Journal of Experimental Biology , 215(1), pp.1-13.
Ibarra-Laclette, E., Lyons, E., Hernández-Guzmán, G., Pérez-Torres, C.A., Carretero-Paulet, L., Chang, T.H., Lan, T., Welch, A.J., Juárez, M.J.A., Simpson, J. and Fernández-Cortés, A., 2013. Architecture and evolution of a minute plant genome. Nature , 498(7452), pp.94-98.
Motta, P.J., 1984. Mechanics and functions of jaw protrusion in teleost fishes: a review. Copeia , pp.1-18.
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Müller, K., Borsch, T., Legendre, L., Porembski, S., Theisen, I. and Barthlott, W., 2004. Evolution of carnivory in Lentibulariaceae and the Lamiales. Plant Biology , 6(4), pp.477-490. 27 Müller, K. and Borsch, T., 2005. Phylogenetics of Utricularia (Lentibulariaceae) and molecular evolution of the trnK intron in a lineage with high substitutional rates. Plant Systematics and Evolution , 250(1-2), pp.39-67.
Roos, G., Van Wassenbergh, S., Herrel, A. and Aerts, P., 2009. Kinematics of suction feeding in the seahorse Hippocampus reidi . Journal of Experimental Biology , 212(21), pp.3490-3498.
Sagaso, A., and Sibaoka, T., 1985. Water extrusion in the trap bladders of Utricularia vulgaris . II. A possible mechanism for water outflow. Bot. Mag. Tokyo , 98, pp.113-124.
Singh, A.K., Prabhakar, S. and Sane, S.P., 2011. The biomechanics of fast prey capture in aquatic bladderworts. Biology letters , p.rsbl20110057.
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Vincent, O., Weißkopf, C., Poppinga, S., Masselter, T., Speck, T., Joyeux, M., Quilliet, C. and Marmottant, P., 2011. Ultra-fast underwater suction traps. Proceedings of the Royal Society of London B: Biological Sciences , 278(1720), pp.2909-2914.
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APPENDICES APPENDIX A: INDEX OF KINEMATICS RECORDINGS 30
The following is a list of videos containing kinematic data for U. gibba in order of date of recording. The time sequence of a suction feeding strike was categorized into eight different visual observations. Time is measured in milliseconds (ms). Observations always begin when the trap door visually starts to open (0.00 ms). Only events that were possible to observe were recorded on the table.
File Name Fluid begins Door is 100% Door begins Prey begins to Prey past Prey past trap Door closed
U. gibba to move open closing move gape door
2010_12_09 n/a 0.400 0.600 0.200 0.400 0.600 11.800
2011_01_27 n/a 3.200 3.200 1.800 2.600 3.200 3.800
2011_02_02 n/a 1.000 13.000 1.000 n/a n/a n/a
2011_02_03 n/a 0.545 0.000 n/a n/a n/a
2011_02_14 n/a 0.545 3.091 0.091 n/a n/a n/a
2011_02_15 n/a 0.636 1.818 0.364 n/a n/a n/a
2011_02_26 n/a 0.000 2.363 18.907
2011_04_06 n/a 0.700 5.400 n/a n/a n/a
2011_04_12 n/a 0.600 4.200 0.200 0.500 0.800
2011_04_13 n/a 0.600 0.200
2011_06_14_002 0.300 0.600 1.400 n/a n/a n/a 7.900
2011_06_14_003 0.000 0.400 n/a n/a n/a
2011_06_15_001 0.200 0.500 n/a n/a n/a 31
File name Fluid begins Door is 100% Door begins Prey begins to Prey past Prey past trap Door closed
U. gibba to move open closing move gape door
2011_06_15_002 0.200 0.700 0.800 n/a n/a n/a 4.300
2011_06_15_003 0.200 0.500 1.200 n/a n/a n/a 5.100
2011_06_15_004 0.300 0.900 n/a n/a n/a
2011_06_16 0.300 0.500 n/a n/a n/a
2011_06_17_002 0.400 0.000 0.500 0.700
2011_06_17_003 0.400 0.000 0.500 0.700
2011_06_29_001 0.100 n/a n/a n/a
2011_06_30_001 0.800 1.500 2.300 n/a n/a n/a 5.300
2011_07_08 1.000 2.600 n/a n/a n/a 6.700 32
File name Fluid begins Door is 100% Door begins Prey begins to Prey past Prey past trap Door closed
U. gibba to move open closing move gape door
2011_07_11_001 0.600 n/a n/a n/a
2011_07_11_002 0.200 0.900 n/a n/a n/a
2011_07_14_002 0.700 n/a n/a n/a
2011_07_20_001 0.200 1.100 1.600 n/a n/a n/a 6.500
2011_07_20_002 0.000 0.600 2.300 n/a n/a n/a 3.300
2011_07_11_002 0.200 0.900 n/a n/a n/a
2011_07_14_002 0.700 n/a n/a n/a
2011_07_20_001 0.200 1.100 1.600 n/a n/a n/a 6.500
2011_07_20_002 0.000 0.600 2.300 n/a n/a n/a 3.300
33
File name Fluid begins Door is 100% Door begins Prey begins to Prey past Prey past trap Door closed
U. gibba to move open closing move gape door
2011_07_20_003 0.000 0.500 1.000 n/a n/a n/a 4.100
2011_07_21 0.700 1.500 3.400 n/a n/a n/a 12.800
2011_07_27 1.000 1.700 0.500 0.900 1.100 3.800
2011_07_28_001 0.200 0.700 1.500 n/a n/a n/a 4.500
2012_12_27_001 0.500 2.860 n/a n/a n/a
2012_12_27_002 0.100 0.340 1.480 n/a n/a n/a 21.040
2014_05_08 0.740 1.000 n/a n/a n/a
2014_05_09_001 0.200 0.340 2.060 n/a n/a n/a 4.620
2014_05_09_002 0.260 0.600 2.260 n/a n/a n/a 5.400
2014_05_10 0.060 0.240 1.800 n/a n/a n/a 6.000 34
File name Fluid begins Door is 100% Door begins Prey begins to Prey past Prey past trap Door closed
U. gibba to move open closing move gape door
2014_05_12_001 -0.080 0.340 2.420 n/a n/a n/a 4.020
2014_05_12_002 -5.820 1.060 1.480 n/a n/a n/a 3.440
2014_05_13 0.080 0.540 1.220 n/a n/a n/a 9.760
2014_05_16 0.160 0.240 n/a n/a n/a
2014_05_26_001 9.980 0.380 n/a n/a n/a
2014_05_26 -0.240 n/a n/a n/a
35
File name Fluid begins Door is 100% Door begins Prey begins to Prey past Prey past trap Door closed
U. vulgaris to move open closing move gape door
2012_10_22 0.550 1.200 n/a n/a n/a 7.250
2012_10_05 0.200 0.550 1.700 n/a n/a n/a 7.900
2012_12_23 1.056 n/a n/a n/a
2012_12_26_001 0.200 n/a n/a n/a 5.700
2012_12_26_002 -0.100 n/a n/a n/a 7.000
2012_12_26_003 0.400 n/a n/a n/a 5.100
2012_12_26_004 0.278 0.778 2.000 n/a n/a n/a 3.889
2012_12_27 0.167 0.944 1.333 n/a n/a n/a 3.500
2012_12_28_001 0.278 0.833 5.167 n/a n/a n/a 9.389
2012_12_28_002 0.056 0.679 2.107 n/a n/a n/a 4.036 36
File name Fluid begins Door is 100% Door begins Prey begins to Prey past Prey past trap Door closed
U. vulgaris to move open closing move gape door
2013_01_15_001 -0.560 0.778 1.333 n/a n/a n/a 4.056
2013_01_15_002 0.167 1.111 1.111 n/a n/a n/a 2.278
2013_01_21_001 -0.464 1.000 3.357 n/a n/a n/a 4.500
2013_01_21_002 0.444 1.222 1.389 n/a n/a n/a
2013_01_23 -0.056 1.278 4.000 n/a n/a n/a 6.222
2013_01_25 0.167 1.111 4.444 n/a n/a n/a 8.389
2013_01_27_001 0.500 1.278 2.833 n/a n/a n/a 5.444
2013_01_27_002 -0.056 0.667 1.444 n/a n/a n/a
2013_01_28 -0.107 1.357 2.607 n/a n/a n/a 5.571
2013_01_29_001 0.000 0.714 3.107 n/a n/a n/a 6.536 37
File name Fluid begins Door is 100% Door begins Prey begins to Prey past Prey past trap Door closed
U. vulgaris to move open closing move gape door
2013_02_01_001 0.889 1.611 2.056 n/a n/a n/a 3.111
2013_02_01_002 0.167 1.056 1.389 n/a n/a n/a 3.333
2013_02_01_003 3.056 3.889 3.944 n/a n/a n/a 5.556
2013_02_01_004 0.333 1.056 n/a n/a n/a
2013_02_01_005 -0.389 0.611 n/a n/a n/a
2013_02_01_006 1.333 1.944 2.389 n/a n/a n/a 7.722
2013_02_02_001 0.333 1.722 2.444 n/a n/a n/a 3.944
2013_02_02_002 0.944 1.500 2.222 n/a n/a n/a 4.000
2013_02_03_001 0.111 0.778 1.278 n/a n/a n/a 3.778
2013_02_03_002 0.389 1.000 3.500 n/a n/a n/a 9.056 38
File name Fluid begins Door is 100% Door begins Prey begins to Prey past Prey past trap Door closed
U. vulgaris to move open closing move gape door
2013_02_03_003 0.111 0.611 1.278 n/a n/a n/a 8.000
2013_02_03_004 0.056 0.889 5.500 n/a n/a n/a 12.667
2013_02_03_005 0.611 0.889 6.611 n/a n/a n/a
2013_02_04_001 -0.056 0.667 1.833 n/a n/a n/a 4.722
2013_02_04_002 0.111 0.778 0.833 n/a n/a n/a 4.000
2013_11_23 0.393 0.500 0.929 0.143 0.786 0.964 5.178
2014_04_17 0.036 0.679 0.964 n/a n/a n/a 3.643
2014_06_03 0.556 0.611 1.500 n/a n/a n/a 8.389
2014_06_05 0.833 1.444 1.556 n/a n/a n/a 3.889
2014_06_06 0.111 0.444 2.056 n/a n/a n/a 6.111 39
File name Fluid begins Door is 100% Door begins Prey begins to Prey past Prey past trap Door closed
U. vulgaris to move open closing move gape door
2014_06_07 0.278 0.611 1.667 n/a n/a n/a 5.722
2014_06_13 0.000 0.667 1.111 0.222 3.056 40
APPENDIX B: INDEX OF FLOW RECORDINGS 42
This appendix lists all recording used for PIV analysis. Each video has its own table listing experimental parameters for each recording: • We use two species, U. gibba and U. vulgaris . • Particles used for PIV were either nylon or luxsil. • The trap was triggered by a whisker, ostracod (=prey), or glass pipette. • Frames per second (FPS) is the speed of the recording. • Resolution differs between videos depending mainly on frame rate. • Traps were recorded from either a front or side view. • ‘Successful’ indicates that there was a response (trap triggered). • Pixel to mm is the calibration factor from pixels to millimeters (mm). • Dimensions measured are gape, bladder length (longest ends), and prey diameter (longest ends). • The top image is a reference shot of that particular recording. Superficially it shows the angle and the overall quality of the recording. The middle image is a graph showing how speed changes in time. A point in the middle of the flow half a gape away from the mouth was chosen to measure this. The y-axis shows speed of particles in meters per second (m/s) and the x-axis shows time in seconds (s). The bottom image is a graph showing how speed changes in space. A transect through the middle of the flow was used to measure this. The y-axis shows speed of particles in meters per second (m/s) and the x-axis shows distance traveled in millimeters (mm). 43
Title: PIV_2011-06-14_002
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Side
Successful Trigger
Pixels per mm 208.9
Gape in Pixels 86.3
Length in Pixels 397.6
Gape in mm 0.413
Length in mm 1.903
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 2.01ms
Time to Peak Speed 0.97ms
Peak Flow Speed 0.16m/s
44
Title: PIV_2011-06-14_003
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Side
Successful Trigger
Pixels per mm 208.9
Gape in Pixels 72.6
Length in Pixels 335.8
Gape in mm 0.348
Length in mm 1.607
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 2.81ms
Time to Peak Speed 1.58ms
Peak Flow Speed 0.21m/s
45
Title: PIV_2011-06-15_001
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Side
Successful Trigger
Pixels per mm 208.9
Gape in Pixels 73.5
Length in Pixels 337.7
Gape in mm 0.352
Length in mm 1.617
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 2.54ms
Time to Peak Speed 1.90ms
Peak Flow Speed 0.17m/s
46
Title: PIV_2011-06-15_002
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View side
Successful Trigger
Pixels per mm 208.9
Gape in Pixels 91.7
Length in Pixels n/a
Gape in mm 0.439
Length in mm n/a
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 1.20ms
Time to Peak Speed 0.63ms
Peak Flow Speed 0.14m/s
47
Title: PIV_2011-06-15_003
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Side
Successful Trigger
Pixels per mm 208.9
Gape in Pixels 80.6
Length in Pixels 322.3
Gape in mm 0.386
Length in mm 1.543
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
48
Title: PIV_2011-06-15_004
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Side
Successful Trigger
Pixels per mm 208.9
Gape in Pixels 70.7
Length in Pixels 339.8
Gape in mm 0.338
Length in mm 1.627
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 2.71ms
Time to Peak Speed 1.38ms
Peak Flow Speed 0.21m/s
49
Title: PIV_2011-06-15_005
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Front
Successful Trigger
Pixels per mm 208.9
Gape in Pixels 87.0
Length in Pixels n/a
Gape in mm 0.416
Length in mm n/a
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 2.04ms
Time to Peak Speed 1.07ms
Peak Flow Speed 0.09m/s
50
Title: PIV_2011-06-16_001
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View side
Successful Trigger
Pixels per mm 208.9
Gape in Pixels 80.5
Length in Pixels 344.4
Gape in mm 0.385
Length in mm 1.649
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 2.61ms
Time to Peak Speed 1.20ms
Peak Flow Speed 0.12m/s
51
Title: PIV_2011-06-16_002
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Front
Successful Trigger
Pixels per mm 208.9
Gape in Pixels
Length in Pixels
Gape in mm
Length in mm
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 2.54ms
Time to Peak Speed 0.71ms
Peak Flow Speed 0.10m/s
52
Title: PIV_2011-06-17_001
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Ostracod
FPS 10,000
Resolution 800x600
View Side
Successful Capture
Pixels per mm 210.8
Gape in Pixels 77.2
Length in Pixels 330.8
Gape in mm 0.366
Length in mm 1.569
Prey Length in Pixels 76.8
Prey Width in Pixels 54.2
Prey Length in mm 0.364
Prey Width in mm 0.257
Flow Event Duration 1.57ms
Time to Peak Speed 1.92ms
Peak Flow Speed 0.16m/s
53
Title: PIV_2011-06-17_002
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Ostracod
FPS 10,000
Resolution 800x600
View Side
Successful Capture
Pixels per mm 210.8
Gape in Pixels 73.1
Length in Pixels 348.8
Gape in mm 0.347
Length in mm 1.655
Prey Length in Pixels 75.76
Prey Width in Pixels 56.4
Prey Length in mm 0.359
Prey Width in mm 0.268
Flow Event Duration 3.21ms
Time to Peak Speed 1.22ms
Peak Flow Speed 0.14m/s
54
Title: PIV_2011-06-17_003
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Ostracod
FPS 10,000
Resolution 800x600
View Side
Successful Capture
Pixels per mm 210.8
Gape in Pixels 68.8
Length in Pixels 351.4
Gape in mm 0.326
Length in mm 1.667
Prey Length in Pixels 62.4
Prey Width in Pixels 43.9
Prey Length in mm 0.296
Prey Width in mm 0.208
Flow Event Duration 2.81ms
Time to Peak Speed 1.38ms
Peak Flow Speed 0.16m/s 55
Title: PIV_2011-06-24_001
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Side
Successful Trigger
Pixels per mm 210.8
Gape in Pixels 93.4
Length in Pixels 389.2
Gape in mm 0.443
Length in mm 1.846
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
56
Title: PIV_2011-06-29_001
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Side
Successful Trigger
Pixels per mm 210.8
Gape in Pixels 67.1
Length in Pixels 331
Gape in mm 0.318
Length in mm 1.570
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 2.01ms
Time to Peak Speed 1.36ms
Peak Flow Speed 0.16m/s
57
Title: PIV_2011-07-01_001
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Side
Successful Trigger
Pixels per mm 210.8
Gape in Pixels 63.8
Length in Pixels 326.2
Gape in mm 0.303
Length in mm 1.547
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 1.40ms
Time to Peak Speed 0.60ms
Peak Flow Speed 0.11m/s
58
Title: PIV_2011-07-08_001
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Front
Successful Trigger
Pixels per mm 210.8
Gape in Pixels 82.3
Length in Pixels 236
Gape in mm 0.390
Length in mm 1.120
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
59
Title: PIV_2011-07-08_002
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Side
Successful Trigger
Pixels per mm 210.8
Gape in Pixels 49.1
Length in Pixels 260.7
Gape in mm 0.233
Length in mm 1.237
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
60
Title: PIV_2011-07-08_003
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Side
Successful Trigger
Pixels per mm 210.8
Gape in Pixels 89.1
Length in Pixels 324.3
Gape in mm 0.423
Length in mm 1.538
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
61
Title: PIV_2011-07-11_001
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Side
Successful Trigger
Pixels per mm 210.8
Gape in Pixels 68.7
Length in Pixels 306.3
Gape in mm 0.326
Length in mm 1.453
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
62
Title: PIV_2011-07-11_002
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Front
Successful Trigger
Pixels per mm 210.8
Gape in Pixels 113.2
Length in Pixels 283
Gape in mm 0.537
Length in mm 1.343
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
63
Title: PIV_2011-07-13_001
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Front
Successful Trigger
Pixels per mm 210.8
Gape in Pixels 111.0
Length in Pixels n/a
Gape in mm 0.527
Length in mm n/a
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 1.74ms
Time to Peak Speed 0.83ms
Peak Flow Speed 0.13m/s
64
Title: PIV_2011-07-14_001
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Side
Successful Trigger
Pixels per mm 210.8
Gape in Pixels 75.6
Length in Pixels 348.2
Gape in mm 0.359
Length in mm 1.652
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
65
Title: PIV_2011-07-14_002
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Side
Successful Trigger
Pixels per mm 210.8
Gape in Pixels 74.2
Length in Pixels 334.2
Gape in mm 0.352
Length in mm 1.585
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed 66
Title: PIV_2011-07-20_001
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Front
Successful Trigger
Pixels per mm 210.8
Gape in Pixels 81.2
Length in Pixels n/a
Gape in mm 0.385
Length in mm n/a
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 2.17ms
Time to Peak Speed 0.94ms
Peak Flow Speed 0.09m/s
67
Title: PIV_2011-07-20_002
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Side
Successful Trigger
Pixels per mm 210.8
Gape in Pixels 80.6
Length in Pixels 305.2
Gape in mm 0.382
Length in mm 1.448
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 2.31ms
Time to Peak Speed 1.03ms
Peak Flow Speed 0.15m/s
68
Title: PIV_2011-07-20_003
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Front
Successful Trigger
Pixels per mm 210.8
Gape in Pixels 64.6
Length in Pixels n/a
Gape in mm 0.306
Length in mm n/a
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 1.71ms
Time to Peak Speed 1.05ms
Peak Flow Speed 0.07m/s
69
Title: PIV_2011-07-20_004
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Front
Successful Trigger
Pixels per mm 210.8
Gape in Pixels 105
Length in Pixels n/a
Gape in mm 0.498
Length in mm n/a
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 2.41ms
Time to Peak Speed 1.20ms
Peak Flow Speed 0.08m/s
70
Title: PIV_2011-07-21_001
Species U. gibba
Illumination PIV
Particles Nylon
Trigger whisker
FPS 10,000
Resolution 800x600
View Front
Successful Trigger
Pixels per mm 210.8
Gape in Pixels 91.8
Length in Pixels n/a
Gape in mm 0.435
Length in mm n/a
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Flow Event Duration 1.74ms
Time to Peak Speed 0.42ms
Peak Flow Speed 0.14m/s
71
Title: PIV_2011-07-21_002
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Front
Successful Trigger
Pixels per mm 210.8
Gape in Pixels 114.2
Length in Pixels 304.2
Gape in mm 0.542
Length in mm 1.443
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
72
Title: PIV_2011-07-21_003
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Side
Successful Trigger
Pixels per mm 210.8
Gape in Pixels 53.5
Length in Pixels 242.1
Gape in mm 0.254
Length in mm 1.148
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
73
Title: PIV_2011-07-27_001
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Ostracod
FPS 10,000
Resolution 800x600
View Side
Successful Capture
Pixels per mm 210.8
Gape in Pixels 68.1
Length in Pixels 248.0
Gape in mm 0.323
Length in mm 1.176
Prey Length in Pixels 56.3
Prey Width in Pixels 44.8
Prey Length in mm 0.267
Prey Width in mm 0.213
Flow Event Duration
Time to Peak Speed
Peak Flow Speed 0.21m/s
74
Title: PIV_2011-07-28_001
Species U. gibba
Illumination PIV
Particles Nylon
Trigger Ostracod
FPS 10,000
Resolution 800x600
View Side
Successful Capture
Pixels per mm 210.8
Gape in Pixels 79.8
Length in Pixels 314.7
Gape in mm 0.379
Length in mm 1.493
Prey Length in Pixels 50.3
Prey Width in Pixels 31.4
Prey Length in mm 0.239
Prey Width in mm 0.149
Flow Event Duration 2.01ms
Time to Peak Speed 1.61ms
Peak Flow Speed 0.12m/s
75
Title: PIV_2012-08-31_001
Species U. gibba
Illumination PIV
Particles Nylon
Trigger whisker
FPS 50,000
Resolution 320x240
View Side
Successful Trigger
Pixels per mm 210.8
Gape in Pixels 44.9
Length in Pixels 232.8
Gape in mm 0.213
Length in mm 1.104
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 1.04ms
Time to Peak Speed 0.44ms
Peak Flow Speed 0.16m/s
76
Title: PIV_2012-12-26_001
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Side
Successful Trigger
Pixels per mm 209.0
Gape in Pixels 140.6
Length in Pixels 543.3
Gape in mm 0.673
Length in mm 2.60
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Flow Event Duration 0.07ms
Time to Peak Speed 1.24ms
Peak Flow Speed 0.14m/s 77
Title: PIV_2012-12-26_002
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 10,000
Resolution 800x600
View Side
Successful Trigger
Pixels per mm 217.0
Gape in Pixels 128.9
Length in Pixels 486.9
Gape in mm 0.594
Length in mm 2.244
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 0.04ms
Time to Peak Speed 1.61ms
Peak Flow Speed 0.12m/s
78
Title: PIV_2012-12-26_004
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 18,000
Resolution 640x480
View Side
Successful Trigger
Pixels per mm 216.0
Gape in Pixels 112.4
Length in Pixels 467.4
Gape in mm 0.520
Length in mm 2.164
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
79
Title: PIV_2012-12-27_001
Species U. gibba
Illumination LED
Particles n/a
Trigger pipette
FPS 50,000
Resolution 320x240
View Side
Successful Trigger
Pixels per mm 210.8
Gape in Pixels 57.5
Length in Pixels n/a
Gape in mm 0.273
Length in mm n/a
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 0.08ms
Time to Peak Speed 0.61ms
Peak Flow Speed 0.20m/s
80
Title: PIV_2012-12-28_001
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 18,000
Resolution 640x480
View Side
Successful Trigger
Pixels per mm 211.1
Gape in Pixels 117.9
Length in Pixels 467.7
Gape in mm 0.559
Length in mm 2.216
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
81
Title: PIV_2012-12-28_002
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 28,000
Resolution 512x384
View Side
Successful Trigger
Pixels per mm 210.0
Gape in Pixels 125.6
Length in Pixels 556.2
Gape in mm 0.598
Length in mm 2.649
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
82
Title: PIV_2013-01-21_002
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 18,000
Resolution 640x480
View Side
Successful Trigger
Pixels per mm 211.0
Gape in Pixels 127.0
Length in Pixels 480.8
Gape in mm 0.602
Length in mm 2.279
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 0.16ms
Time to Peak Speed 0.93ms
Peak Flow Speed 0.38m/s
83
Title: PIV_2013-01-23_001
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 18,000
Resolution 640x480
View Side
Successful Trigger
Pixels per mm 211.1
Gape in Pixels 129.4
Length in Pixels 518.5
Gape in mm 0.613
Length in mm 2.456
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
84
Title: PIV_2013-01-25_001
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 18,000
Resolution 640x480
View Side
Successful Trigger
Pixels per mm 209.0
Gape in Pixels 79.4
Length in Pixels 347.2
Gape in mm 0.380
Length in mm 1.661
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
85
Title: PIV_2013-02-01_001
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 18,000
Resolution 640x480
View Side
Successful Trigger
Pixels per mm 213.0
Gape in Pixels 133.3
Length in Pixels 610.9
Gape in mm 0.626
Length in mm 2.868
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 0.02ms
Time to Peak Speed 1.16ms
Peak Flow Speed 0.15m/s
86
Title: PIV_2013-02-01_002
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 18,000
Resolution 640x480
View Side
Successful Trigger
Pixels per mm 209.0
Gape in Pixels 138.9
Length in Pixels 534.2
Gape in mm 0.665
Length in mm 2.556
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 0.07ms
Time to Peak Speed 0.87ms
Peak Flow Speed 0.36m/s
87
Title: PIV_2013-02-01_003
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 18,000
Resolution 640x480
View Side
Successful Trigger
Pixels per mm 210.0
Gape in Pixels 132.4
Length in Pixels 544.6
Gape in mm 0.630
Length in mm 2.593
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed 88
Title: PIV_2013-02-01_004
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 18,000
Resolution 640x480
View Side
Successful Trigger
Pixels per mm 208.0
Gape in Pixels 104.6
Length in Pixels 470.2
Gape in mm 0.503
Length in mm 2.261
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 0.10ms
Time to Peak Speed 1.19ms
Peak Flow Speed 0.30m/s
89
Title: PIV_2013-02-01_006
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 18,000
Resolution 640x480
View Side
Successful Trigger
Pixels per mm 210.0
Gape in Pixels 127.3
Length in Pixels 510.8
Gape in mm 0.606
Length in mm 2.432
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 0.13ms
Time to Peak Speed 0.43ms
Peak Flow Speed 0.28m/s
90
Title: PIV_2013-02-02_001
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 18,000
Resolution 640x480
View Side
Successful Trigger
Pixels per mm 214.0
Gape in Pixels 127.7
Length in Pixels 531.0
Gape in mm 0.597
Length in mm 2.481
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
91
Title: PIV_2013-02-02_002
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 18,000
Resolution 640x480
View Side
Successful Trigger
Pixels per mm 215.0
Gape in Pixels 138.1
Length in Pixels 539.5
Gape in mm 0.642
Length in mm 2.509
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
92
Title: PIV_2013-02-03_003
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 18,000
Resolution 640x480
View Side
Successful Trigger
Pixels per mm 213.0
Gape in Pixels 129.4
Length in Pixels 658.3
Gape in mm 0.608
Length in mm 3.091
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
93
Title: PIV_2013-02-03_004
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 18,000
Resolution 640x480
View Side
Successful Trigger
Pixels per mm 211.1
Gape in Pixels 134.5
Length in Pixels 536.1
Gape in mm 0.637
Length in mm 2.540
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 0.11ms
Time to Peak Speed 2.62ms
Peak Flow Speed 0.26m/s
94
Title: PIV_2013-02-03_005
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 18,000
Resolution 640x480
View side
Successful Trigger
Pixels per mm 214.1
Gape in Pixels 128.2
Length in Pixels 545.6
Gape in mm 0.599
Length in mm 2.548
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
95
Title: PIV_2013-02-04_001
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 18,000
Resolution 640x480
View Side
Successful Trigger
Pixels per mm 210.0
Gape in Pixels 141.0
Length in Pixels 544.0
Gape in mm 0.671
Length in mm 2.590
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Flow Event Duration 0.05ms
Time to Peak Speed 1.19ms
Peak Flow Speed 0.24m/s
96
Title: PIV_2013-02-04_002
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Whisker
FPS 18,000
Resolution 640x480
View Side
Successful Trigger
Pixels per mm 212.0
Gape in Pixels 150.4
Length in Pixels 546.6
Gape in mm 0.709
Length in mm 2.578
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 0.02ms
Time to Peak Speed 0.56ms
Peak Flow Speed 0.14m/s
97
Title: PIV_2013-11-23_001
Species U. vulgaris
Illumination PIV
Particles Nylon
Trigger Ostracod
FPS 28,001
Resolution 512x384
View side
Successful Capture
Pixels per mm 102.0
Gape in Pixels 55.0
Length in Pixels 189.5
Gape in mm 0.539
Length in mm 1.858
Prey Length in Pixels 38.8
Prey Width in Pixels 24.8
Prey Length in mm 0.380
Prey Width in mm 0.243
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
98
Title: PIV_2014-04-17_001
Species U. vulgaris
Illumination PIV
Particles Luxsil
Trigger Whisker
FPS 28,001
Resolution 512x384
View Side
Successful Trigger
Pixels per mm 102.0
Gape in Pixels 54.6
Length in Pixels 248.0
Gape in mm 0.535
Length in mm 2.431
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed 99
Title: PIV_2014-05-13_001
Species U. gibba
Illumination PIV
Particles Luxsil
Trigger Whisker
FPS 50,000
Resolution 320x240
View Side
Successful Triggered
Pixels per mm 100.0
Gape in Pixels 50.2
Length in Pixels 182.1
Gape in mm 0.502
Length in mm 1.821
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration 1.97ms
Time to Peak Speed 1.04ms
Peak Flow Speed 0.23m/s
100
Title: PIV_2014-06-05_001
Species U. vulgaris
Illumination PIV
Particles Luxsil
Trigger Whisker
FPS 18,000
Resolution 640x480
View side
Successful Triggered
Pixels per mm 218.3
Gape in Pixels 116.7
Length in Pixels 477.3
Gape in mm 0.535
Length in mm 2.186
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
101
Title: PIV_2014-06-06_001
Species U. vulgaris
Illumination PIV
Particles Luxsil
Trigger Whisker
FPS 18,000
Resolution 640x480
View front
Successful Triggered
Pixels per mm 212.3
Gape in Pixels 198.4
Length in Pixels 477.3
Gape in mm 0.935
Length in mm 2.248
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
102
Title: PIV_2014-06-07_001
Species U. vulgaris
Illumination PIV
Particles Luxsil
Trigger Whisker
FPS 18,000
Resolution 640x480
View Front
Successful Triggered
Pixels per mm 210.1
Gape in Pixels 192.4
Length in Pixels 429.0
Gape in mm 0.916
Length in mm 2.042
Prey Length in Pixels n/a
Prey Width in Pixels n/a
Prey Length in mm n/a
Prey Width in mm n/a
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
103
Title: PIV_2014-06-13_001
Species U. vulgaris
Illumination PIV
Particles Luxsil
Trigger Ostracod
FPS 18,000
Resolution 640x480
View Side
Successful Missed
Pixels per mm 216.3
Gape in Pixels 136.2
Length in Pixels 429.0
Gape in mm 0.630
Length in mm 1.983
Prey Length in Pixels 117.6
Prey Width in Pixels 70.6
Prey Length in mm 0.5437
Flow Event Duration
Time to Peak Speed
Peak Flow Speed
Fresno State
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Matthew D. Brown
Type full name as it appears on submission
April 18, 2016
Date