Colin Sean

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Colin

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Sean

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0000-0003-4463-5588

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Neuromechanical wave resonance in jellyfish swimming

(National Academy of Sciences, 2021-03-16) Hoover, Alexander P. ; Xu, Nicole W. ; Gemmell, Brad J. ; Colin, Sean P. ; Costello, John H. ; Dabiri, John O. ; Miller, Laura A.

For organisms to have robust locomotion, their neuromuscular organization must adapt to constantly changing environments. In jellyfish, swimming robustness emerges when marginal pacemakers fire action potentials throughout the bell’s motor nerve net, which signals the musculature to contract. The speed of the muscle activation wave is dictated by the passage times of the action potentials. However, passive elastic material properties also influence the emergent kinematics, with time scales independent of neuromuscular organization. In this multimodal study, we examine the interplay between these two time scales during turning. A three-dimensional computational fluid–structure interaction model of a jellyfish was developed to determine the resulting emergent kinematics, using bidirectional muscular activation waves to actuate the bell rim. Activation wave speeds near the material wave speed yielded successful turns, with a 76-fold difference in turning rate between the best and worst performers. Hyperextension of the margin occurred only at activation wave speeds near the material wave speed, suggesting resonance. This hyperextension resulted in a 34-fold asymmetry in the circulation of the vortex ring between the inside and outside of the turn. Experimental recording of the activation speed confirmed that jellyfish actuate within this range, and flow visualization using particle image velocimetry validated the corresponding fluid dynamics of the numerical model. This suggests that neuromechanical wave resonance plays an important role in the robustness of an organism’s locomotory system and presents an undiscovered constraint on the evolution of flexible organisms. Understanding these dynamics is essential for developing actuators in soft body robotics and bioengineered pumps.
Swimming kinematics and performance of spinal transected lampreys with different levels of axon regeneration

(The Company of Biologists, 2021-11-05) Fies, Jacob ; Gemmell, Brad J. ; Fogerson, Stephanie M. ; Morgan, Jennifer R. ; Tytell, Eric D. ; Colin, Sean P.

Axon regeneration is critical for restoring neural function after spinal cord injury. This has prompted a series of studies on the neural and functional recovery of lampreys after spinal cord transection. Despite this, there are still many basic questions remaining about how much functional recovery depends on axon regeneration. Our goal was to examine how swimming performance is related to degree of axon regeneration in lampreys recovering from spinal cord transection by quantifying the relationship between swimming performance and percent axon regeneration of transected lampreys after 11 weeks of recovery. We found that while swimming speeds varied, they did not relate to percent axon regeneration. In fact, swimming speeds were highly variable within individuals, meaning that most individuals could swim at both moderate and slow speeds, regardless of percent axon regeneration. However, none of the transected individuals were able to swim as fast as the control lampreys. To swim fast, control lampreys generated high amplitude body waves with long wavelengths. Transected lampreys generated body waves with lower amplitude and shorter wavelengths than controls, and to compensate, transected lampreys increased their wave frequencies to swim faster. As a result, transected lampreys had significantly higher frequencies than control lampreys at comparable swimming velocities. These data suggest that the control lampreys swam more efficiently than transected lampreys. In conclusion, there appears to be a minimal recovery threshold in terms of percent axon regeneration required for lampreys to be capable of swimming; however, there also seems to be a limit to how much they can behaviorally recover.
An algorithm to estimate unsteady and quasi-steady pressure fields from velocity field measurements

(Company of Biologists, 2013-10-10) Dabiri, John O. ; Bose, Sanjeeb ; Gemmell, Brad J. ; Colin, Sean P. ; Costello, John H.

We describe and characterize a method for estimating the pressure field corresponding to velocity field measurements such as those obtained by using particle image velocimetry. The pressure gradient is estimated from a time series of velocity fields for unsteady calculations or from a single velocity field for quasi-steady calculations. The corresponding pressure field is determined based on median polling of several integration paths through the pressure gradient field in order to reduce the effect of measurement errors that accumulate along individual integration paths. Integration paths are restricted to the nodes of the measured velocity field, thereby eliminating the need for measurement interpolation during this step and significantly reducing the computational cost of the algorithm relative to previous approaches. The method is validated by using numerically simulated flow past a stationary, two-dimensional bluff body and a computational model of a three-dimensional, self-propelled anguilliform swimmer to study the effects of spatial and temporal resolution, domain size, signal-to-noise ratio and out-of-plane effects. Particle image velocimetry measurements of a freely swimming jellyfish medusa and a freely swimming lamprey are analyzed using the method to demonstrate the efficacy of the approach when applied to empirical data.
Propulsive design principles in a multi-jet siphonophore.

(Company of Biologists, 2019-02-27) Sutherland, Kelly R. ; Gemmell, Brad J. ; Colin, Sean P. ; Costello, John H.

Coordination of multiple propulsors can provide performance benefits in swimming organisms. Siphonophores are marine colonial organisms that orchestrate the motion of multiple swimming zooids for effective swimming. However, the kinematics at the level of individual swimming zooids (nectophores) have not been examined in detail. We used high speed, high resolution microvideography and particle image velocimetry (PIV) of the physonect siphonophore, Nanomia bijuga, to study the motion of the nectophores and the associated fluid motion during jetting and refilling. The integration of nectophore and velum kinematics allow for a high-speed (maximum ∼1 m s−1), narrow (1-2 mm) jet and rapid refill as well as a 1:1 ratio of jetting to refill time. Scaled to the 3 mm nectophore length, jet speeds reach >300 lengths s−1. Overall swimming performance is enhanced by velocity gradients produced in the nectophore during refill, which lead to a high pressure region that produces forward thrust. Generating thrust during both the jet and refill phases augments the distance travelled by 17% over theoretical animals, which generate thrust only during the jet phase. The details of velum kinematics and associated fluid mechanics elucidate how siphonophores effectively navigate three-dimensional space and could be applied to exit flow parameters in multijet underwater vehicles.
Danger of zooplankton feeding : the fluid signal generated by ambush-feeding copepods

( 2010-06) Kiørboe, Thomas ; Jiang, Houshuo ; Colin, Sean P.

Zooplankton feed in either of three ways: they generate a feeding current, cruise through the water, or they are ambush feeders. Each mode generates different hydrodynamic disturbances and hence exposes the grazers differently to mechanosensory predators. Ambush feeders sink slowly and therefore perform occasional upward repositioning jumps. We quantified the fluid disturbance generated by repositioning jumps in a mm-sized copepod (Re ~ 40). The kick of the swimming legs generates a viscous vortex ring in the wake; another ring of similar intensity but opposite rotation is formed around the decelerating copepod. A simple analytical model, that of an impulsive point force, properly describes the observed flow field as a function of the momentum of the copepod, including the translation of the vortex and its spatial extension and temporal decay. We show that the time-averaged fluid signal and the consequent predation risk is much less for an ambush feeding than a cruising or hovering copepod for small individuals, while the reverse is true for individuals larger than about 1 mm. This makes inefficient ambush feeding feasible in small copepods and is consistent with the observation that ambush feeding copepods in the ocean are all small, while larger species invariably use hovering or cruising feeding strategies.
Passive energy recapture in jellyfish contributes to propulsive advantage over other metazoans

( 2013-08) Gemmell, Brad J. ; Costello, John H. ; Colin, Sean P. ; Stewart, Colin J. ; Dabiri, John O. ; Tafti, Danesh ; Priya, Shashank

Gelatinous zooplankton populations are well known for their ability to take over perturbed ecosystems. The ability of these animals to outcompete and functionally replace fish that exhibit an effective visual predatory mode is counterintuitive because jellyfish are described as inefficient swimmers that must rely on direct contact with prey in order to feed. We show that jellyfish exhibit a novel mechanism of passive energy recapture, which is exploited to allow jellyfish to travel 30 percent further each swimming cycle, thereby reducing metabolic energy demand by swimming muscles. By accounting for large interspecific differences in net metabolic rates, we demonstrate, contrary to prevailing views, the jellyfish (Aurelia aurita) is one of the most energetically efficient propulsors on the planet, exhibiting a cost-of-transport (J kg-1 m-1) lower than other metazoans. We estimate that reduced metabolic demand by passive energy recapture improves cost-of-transport by 48%, allowing jellyfish to achieve the large sizes required for sufficient prey encounters. Pressure calculations, using both computational fluid dynamics (CFD) and a new method from empirical velocity field measurements demonstrate that this extra thrust results from positive pressure created by a vortex ring underneath the bell during the refilling phase of swimming. These results demonstrate a physical basis for the ecological success of medusan swimmers despite their simple body plan. Results from this study also have implications for bio-inspired design where low-energy propulsion is required.
Thrust generation during steady swimming and acceleration from rest in anguilliform swimmers

(Company of Biologists, 2019-11-18) Du Clos, Kevin T. ; Dabiri, John O. ; Costello, John H. ; Colin, Sean P. ; Morgan, Jennifer R. ; Fogerson, Stephanie M. ; Gemmell, Brad J.

Escape swimming is a crucial behavior by which undulatory swimmers evade potential threats. The hydrodynamics of escape swimming have not been well studied, particularly for anguilliform swimmers, such as the sea lamprey Petromyzon marinus. For this study, we compared the kinematics and hydrodynamics of larval sea lampreys with those of lampreys accelerating from rest during escape swimming. We used experimentally derived velocity fields to calculate pressure fields and distributions of thrust and drag along the body. Lampreys initiated acceleration from rest with the formation of a high-amplitude body bend at approximately one-quarter body length posterior to the head. This deep body bend produced two high-pressure regions from which the majority of thrust for acceleration was derived. In contrast, steady swimming was characterized by shallower body bends and negative-pressure-derived thrust, which was strongest near the tail. The distinct mechanisms used for steady swimming and acceleration from rest may reflect the differing demands of the two behaviors. High-pressure-based mechanisms, such as the one used for acceleration from rest, could also be important for low-speed maneuvering during which drag-based turning mechanisms are less effective. The design of swimming robots may benefit from the incorporation of such insights from unsteady swimming.
Fluid interactions that enable stealth predation by the upstream-foraging hydromedusa Craspedacusta sowerbyi

(Marine Biological Laboratory, 2013-09-01) Lucas, Kelsey N. ; Colin, Sean P. ; Costello, John H. ; Katija, Kakani ; Klos, E.

Unlike most medusae that forage with tentacles trailing behind their bells, several species forage upstream of their bells using aborally located tentacles. It has been hypothesized that these medusae forage as stealth predators by placing their tentacles in more quiescent regions of flow around their bells. Consequently, they are able to capture highly mobile, sensitive prey. We used digital particle image velocimetry (DPIV) to quantitatively characterize the flow field around Craspedacusta sowerbyi, a freshwater upstream-foraging hydromedusa, to evaluate the mechanics of its stealth predation. We found that fluid velocities were minimal in front and along the sides of the bell where the tentacles are located. As a result, the deformation rates in the regions where the tentacles are located were low, below the threshold rates required to elicit an escape response in several species of copepods. Estimates of their encounter volume rates were examined on the basis of flow past the tentacles, and trade-offs associated with tentacle characteristics were evaluated.
Maneuvering performance in the colonial siphonophore, Nanomia bijuga

(MDPI, 2019-09-05) Sutherland, Kelly R. ; Gemmell, Brad J. ; Colin, Sean P. ; Costello, John H.

The colonial cnidarian, Nanomia bijuga, is highly proficient at moving in three-dimensional space through forward swimming, reverse swimming and turning. We used high speed videography, particle tracking, and particle image velocimetry (PIV) with frame rates up to 6400 s−1 to study the kinematics and fluid mechanics of N. bijuga during turning and reversing. N. bijuga achieved turns with high maneuverability (mean length–specific turning radius, R/L = 0.15 ± 0.10) and agility (mean angular velocity, ω = 104 ± 41 deg. s−1). The maximum angular velocity of N. bijuga, 215 deg. s−1, exceeded that of many vertebrates with more complex body forms and neurocircuitry. Through the combination of rapid nectophore contraction and velum modulation, N. bijuga generated high speed, narrow jets (maximum = 1063 ± 176 mm s−1; 295 nectophore lengths s−1) and thrust vectoring, which enabled high speed reverse swimming (maximum = 134 ± 28 mm s−1; 37 nectophore lengths s−1) that matched previously reported forward swimming speeds. A 1:1 ratio of forward to reverse swimming speed has not been recorded in other swimming organisms. Taken together, the colonial architecture, simple neurocircuitry, and tightly controlled pulsed jets by N. bijuga allow for a diverse repertoire of movements. Considering the further advantages of scalability and redundancy in colonies, N. bijuga is a model system for informing underwater propulsion and navigation of complex environments.
How the bending kinematics of swimming lampreys build negative pressure fields for suction thrust

(Company of Biologists, 2016-12-14) Gemmell, Brad J. ; Fogerson, Stephanie M. ; Costello, John H. ; Morgan, Jennifer R. ; Dabiri, John O. ; Colin, Sean P.

Swimming animals commonly bend their bodies to generate thrust. For undulating animals such as eels and lampreys, their bodies bend in the form of waves that travel from head to tail. These kinematics accelerate the flow of adjacent fluids, which alters the pressure field in a manner that generates thrust. We used a comparative approach to evaluate the cause-and-effect relationships in this process by quantifying the hydrodynamic effects of body kinematics at the body–fluid interface of the lamprey, Petromyzon marinus, during steady-state swimming. We compared the kinematics and hydrodynamics of healthy control lampreys to lampreys whose spinal cord had been transected mid-body, resulting in passive kinematics along the posterior half of their body. Using high-speed particle image velocimetry (PIV) and a method for quantifying pressure fields, we detail how the active bending kinematics of the control lampreys were crucial for setting up strong negative pressure fields (relative to ambient fields) that generated high-thrust regions at the bends as they traveled all along the body. The passive kinematics of the transected lamprey were only able to generate significant thrust at the tail, relying on positive pressure fields. These different pressure and thrust scenarios are due to differences in how active versus passive body waves generated and controlled vorticity. This demonstrates why it is more effective for undulating lampreys to pull, rather than push, themselves through the fluid.
Propulsion in cubomedusae : mechanisms and utility

(Public Library of Science, 2013-02-20) Colin, Sean P. ; Costello, John H. ; Katija, Kakani ; Seymour, Jamie ; Kiefer, Kristen

Evolutionary constraints which limit the forces produced during bell contractions of medusae affect the overall medusan morphospace such that jet propulsion is limited to only small medusae. Cubomedusae, which often possess large prolate bells and are thought to swim via jet propulsion, appear to violate the theoretical constraints which determine the medusan morphospace. To examine propulsion by cubomedusae, we quantified size related changes in wake dynamics, bell shape, swimming and turning kinematics of two species of cubomedusae, Chironex fleckeri and Chiropsella bronzie. During growth, these cubomedusae transitioned from using jet propulsion at smaller sizes to a rowing-jetting hybrid mode of propulsion at larger sizes. Simple modifications in the flexibility and kinematics of their velarium appeared to be sufficient to alter their propulsive mode. Turning occurs during both bell contraction and expansion and is achieved by generating asymmetric vortex structures during both stages of the swimming cycle. Swimming characteristics were considered in conjunction with the unique foraging strategy used by cubomedusae.
A ctenophore (comb jelly) employs vortex rebound dynamics and outperforms other gelatinous swimmers

(The Royal Society, 2019-03-20) Gemmell, Brad J. ; Colin, Sean P. ; Costello, John H. ; Sutherland, Kelly R.

Gelatinous zooplankton exhibit a wide range of propulsive swimming modes. One of the most energetically efficient is the rowing behaviour exhibited by many species of schyphomedusae, which employ vortex interactions to achieve this result. Ctenophores (comb jellies) typically use a slow swimming, cilia-based mode of propulsion. However, species within the genus Ocyropsis have developed an additional propulsive strategy of rowing the lobes, which are normally used for feeding, in order to rapidly escape from predators. In this study, we used high-speed digital particle image velocimetry to examine the kinematics and fluid dynamics of this rarely studied propulsive mechanism. This mechanism allows Ocyropsis to achieve size-adjusted speeds that are nearly double those of other large gelatinous swimmers. The investigation of the fluid dynamic basis of this escape mode reveals novel vortex interactions that have not previously been described for other biological propulsion systems. The arrangement of vortices during escape swimming produces a similar configuration and impact as that of the well-studied ‘vortex rebound’ phenomenon which occurs when a vortex ring approaches a solid wall. These results extend our understanding of how animals use vortex–vortex interactions and provide important insights that can inform the bioinspired engineering of propulsion systems.
Raw capture efficiency data of scyphomedusae from video analysis collected in Woods Hole, MA beginning in 2015.

(Biological and Chemical Oceanography Data Management Office (BCO-DMO). Contact: bco-dmo-data@whoi.edu, 2019-04-04) Colin, Sean

Raw capture efficiency data of scyphomedusae from video analysis collected in Woods Hole, MA beginning in 2015. For a complete list of measurements, refer to the full dataset description in the supplemental file 'Dataset_description.pdf'. The most current version of this dataset is available at: https://www.bco-dmo.org/dataset/683750
Oceanic lobate ctenophores possess feeding mechanics similar to the impactful coastal species Mnemiopsis leidyi

(Association for the Sciences of Limnology and Oceanography, 2022-09-23) Cordeiro, Malaika ; Costello, John H. ; Gemmell, Brad J. ; Sutherland, Kelly R. ; Colin, Sean P.

Lobate ctenophores are often numerically dominant members of oceanic epipelagic and midwater ecosystems. Despite this, little is known about their trophic ecology. Multiple, co‐occurring species are often found in these ecosystems and appear to feed similarly via feeding currents that entrain prey. We quantified the hydrodynamics, morphology, and behavior of four co‐occurring, cosmopolitan lobate species (Eurhamphaea vexilligera, Ocyropsis crystallina, Bolinopsis vitrea, and Leucothea multicornis) to evaluate whether their feeding mechanics lead to differential feeding rates and prey selection. We compared the feeding characteristics of these four oceanic species to the coastal lobate ctenophore, Mnemiopsis leidyi, which is known as a voracious zooplanktivore. We found that despite their morphological diversity, the five lobate species used the same mechanism to generate their feeding current—the hydrodynamics of their feeding currents were similarly laminar and with very low fluid deformation rates. Despite having similar feeding current traits, the species had different in situ swimming behaviors and feeding postures. We show that these different behaviors and postures lead to different prey encounter rates and that several of the oceanic species have the potential to feed at rates similar to or greater than M. leidyi. As such, the individual and combined trophic impact of oceanic lobate ctenophores is likely to be much greater than previously predicted.
Suction-based propulsion as a basis for efficient animal swimming

(Nature Publishing Group, 2015-11-03) Gemmell, Brad J. ; Colin, Sean P. ; Costello, John H. ; Dabiri, John O.

A central and long-standing tenet in the conceptualization of animal swimming is the idea that propulsive thrust is generated by pushing the surrounding water rearward. Inherent in this perspective is the assumption that locomotion involves the generation of locally elevated pressures in the fluid to achieve the expected downstream push of the surrounding water mass. Here we show that rather than pushing against the surrounding fluid, efficient swimming animals primarily pull themselves through the water via suction. This distinction is manifested in dominant low-pressure regions generated in the fluid surrounding the animal body, which are observed by using particle image velocimetry and a pressure calculation algorithm applied to freely swimming lampreys and jellyfish. These results suggest a rethinking of the evolutionary adaptations observed in swimming animals as well as the mechanistic basis for bio-inspired and biomimetic engineered vehicles.
Prey capture by the cosmopolitan hydromedusae, Obelia spp., in the viscous regime

(John Wiley & Sons, 2016-08-29) Sutherland, Kelly R. ; Gemmell, Brad J. ; Colin, Sean P. ; Costello, John H.

Obelia spp. are cnidarian hydromedusae with a cosmopolitan distribution but very little is known about their feeding. The small size of Obelia (bell diameter ∼ 1 mm, tentacle width ∼ 0.05 mm) suggests that feeding occurs in a viscous regime characterized by thick boundary layers. During feeding observations with a natural prey assemblage the majority of prey were captured at the tentacle tips during the contraction phase. Swimming kinematics from high speed videography confirmed that swimming was a low Re number process (Re < 50) and showed that maximum tentacle velocities occurred at the tentacle tips midway through a bell contraction. Flow visualizations from particle image velocimetry demonstrated that fluid motion between the tentacles was limited and that velocities were highest at the tentacle tips, leading to a thinning of boundary layer in this region. The highest nematocyst densities were observed in this same region of the tentacle tips. Taken together, the body kinematics, flow visualizations and nematocyst distributions of Obelia explain how these predators are able to shed viscous boundary layers to effectively capture microplanktonic prey. Our findings help explain how other small feeding-current medusae whose feeding interactions are governed by viscosity are able to successfully forage.
Widespread utilization of passive energy recapture in swimming medusae

(The Company of Biologists, 2017-11-27) Gemmell, Brad J. ; Colin, Sean P. ; Costello, John H.

Recently, it has been shown that some medusae are capable of swimming very efficiently, i.e. with a low cost of transport, and that this is in part due to passive energy recapture (PER) which occurs during bell relaxation. We compared the swimming kinematics among a diverse array of medusae, varying in taxonomy, morphology and propulsive and foraging modes, in order to evaluate the prevalence of PER in medusae. We found that while PER was common among taxa, the magnitude of the contribution to overall swimming varied greatly. The ability of medusae to utilize PER was not related to morphology and swimming performance but was controlled by their swimming kinematics. Utilizing PER required the medusae to pause after bell expansion and individuals could modulate their PER by changing their pause duration. PER can greatly enhance swimming efficiency but there appear to be trade-offs associated with utilizing PER.
Multi-jet propulsion organized by clonal development in a colonial siphonophore

(Nature Publishing Group, 2015-09-01) Costello, John H. ; Colin, Sean P. ; Gemmell, Brad J. ; Dabiri, John O. ; Sutherland, Kelly R.

Physonect siphonophores are colonial cnidarians that are pervasive predators in many neritic and oceanic ecosystems. Physonects employ multiple, clonal medusan individuals, termed nectophores, to propel an aggregate colony. Here we show that developmental differences between clonal nectophores of the physonect Nanomia bijuga produce a division of labour in thrust and torque production that controls direction and magnitude of whole-colony swimming. Although smaller and less powerful, the position of young nectophores near the apex of the nectosome allows them to dominate torque production for turning, whereas older, larger and more powerful individuals near the base of the nectosome contribute predominantly to forward thrust production. The patterns we describe offer insight into the biomechanical success of an ecologically important and widespread colonial animal group, but, more broadly, provide basic physical understanding of a natural solution to multi-engine organization that may contribute to the expanding field of underwater-distributed propulsion vehicle design.
Hydrodynamics of vortex generation during bell contraction by the hydromedusa Eutonina indicans (Romanes, 1876).

(MDPI, 2019-07-05) Costello, John H. ; Colin, Sean P. ; Gemmell, Brad J. ; Dabiri, John O.

Swimming bell kinematics and hydrodynamic wake structures were documented during multiple pulsation cycles of a Eutonina indicans (Romanes, 1876) medusa swimming in a predominantly linear path. Bell contractions produced pairs of vortex rings with opposite rotational sense. Analyses of the momentum flux in these wake structures demonstrated that vortex dynamics related directly to variations in the medusa swimming speed. Furthermore, a bulk of the momentum flux in the wake was concentrated spatially at the interfaces between oppositely rotating vortices rings. Similar thrust-producing wake structures have been described in models of fish swimming, which posit vortex rings as vehicles for energy transport from locations of body bending to regions where interacting pairs of opposite-sign vortex rings accelerate the flow into linear propulsive jets. These findings support efforts toward soft robotic biomimetic propulsion
Field testing of biohybrid robotic jellyfish to demonstrate enhanced swimming speeds

(MDPI, 2020-11-21) Xu, Nicole W. ; Townsend, James P. ; Costello, John H. ; Colin, Sean P. ; Gemmell, Brad J. ; Dabiri, John O.

Biohybrid robotic designs incorporating live animals and self-contained microelectronic systems can leverage the animals’ own metabolism to reduce power constraints and act as natural chassis and actuators with damage tolerance. Previous work established that biohybrid robotic jellyfish can exhibit enhanced speeds up to 2.8 times their baseline behavior in laboratory environments. However, it remains unknown if the results could be applied in natural, dynamic ocean environments and what factors can contribute to large animal variability. Deploying this system in the coastal waters of Massachusetts, we validate and extend prior laboratory work by demonstrating increases in jellyfish swimming speeds up to 2.3 times greater than their baseline, with absolute swimming speeds up to 6.6 ± 0.3 cm s−1. These experimental swimming speeds are predicted using a hydrodynamic model with morphological and time-dependent input parameters obtained from field experiment videos. The theoretical model can provide a basis to choose specific jellyfish with desirable traits to maximize enhancements from robotic manipulation. With future work to increase maneuverability and incorporate sensors, biohybrid robotic jellyfish can potentially be used to track environmental changes in applications for ocean monitoring.

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