Wheeler Jeanette D.

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Jeanette D.

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Now showing 1 - 7 of 7
  • Preprint
    Desperate planktotrophs : decreased settlement selectivity with age in competent eastern oyster Crassostrea virginica larvae
    ( 2018-07) Meyer, Kirstin S. ; Wheeler, Jeanette D. ; Houlihan, Erin ; Mullineaux, Lauren S.
    For larvae of benthic marine invertebrate species, settlement from planktonic to benthic life is a critical transition. The “desperate larva” concept describes the tendency of larvae to accept suboptimal settlement habitats as they age. We quantified swimming behavior in planktotrophic larvae of the eastern oyster, Crassostrea virginica, to determine whether settlement behaviors, such as swimming downward and remaining on the bottom, increased with age and whether these ontogenetic changes were more apparent in larvae exposed to suboptimal conditions than to preferred conditions (settlement cue absent or present, respectively). In two experiments, the proportion of competent larvae remaining near the bottom of experimental flasks (indicating settlement) increased with larval age, but only in larvae that were not exposed to the settlement cue. This result is consistent with the hypothesis that larvae encountering suboptimal habitat become “desperate” (i.e. more likely to settle) as they age. Exploratory behaviors, such as upward swimming, meandering, or helices, were expected to decrease with age, especially in the absence of the settlement cue, but this pattern was detected in only one of the five swimming metrics tested (helices in downward swimming larvae). Surprisingly, pre-competent larvae exhibited settlement behavior when exposed to the cue, raising the question of whether a response at this stage would have positive or negative consequences. Acceptance of suboptimal settlement habitats by aging larvae may increase the resilience of a species by allowing populations to persist in variable environmental conditions.
  • Thesis
    Behavioral responses of invertebrate larvae to water column cues
    (Massachusetts Institute of Technology and Woods Hole Oceanographic Institution, 2016-02) Wheeler, Jeanette D.
    Many benthic marine invertebrates have two-phase life histories, relying on planktonic larval stages for dispersal and exchange of individuals between adult populations. Historically, larvae were considered passive drifters in prevailing ocean currents. More recently, however, the paradigm has shifted toward active larval behavior mediating transport in the water column. Larvae in the plankton encounter a variety of physical, chemical, and biological cues, and their behavioral responses to these cues may directly impact transport, survival, settlement, and successful recruitment. In this thesis, I investigated the effects of turbulence, light, and conspecific adult exudates on larval swimming behavior. I focused on two invertebrate species of distinct morphologies: the purple urchin Arbacia punctulata, which was studied in pre-settlement planktonic stages, and the Eastern oyster Crassostrea virginica, which was studied in the competent-to-settle larval stage. From this work, I developed a conceptual framework within which larval behavior is understood as being driven simultaneously by external environmental cues and by larval age. As no a priori theory for larval behavior is derivable from first principles, it is only through experimental work that we are able to access behaviors and tie them back to specific environmental triggers. In this work, I studied the behavioral responses of larvae at the individual level, but those dynamics are likely playing out at larger scales in the ocean, impacting population connectivity, community structure, and resilience. In this way, my work represents progress in understanding how the ocean environment and larval behavior couple to influence marine ecological processes.
  • Article
    Ontogenetic changes in larval swimming and orientation of pre-competent sea urchin Arbacia punctulata in turbulence
    (Company of Biologists, 2016-05-04) Wheeler, Jeanette D. ; Chan, Kit Yu Karen ; Anderson, Erik J. ; Mullineaux, Lauren S.
    Many marine organisms have complex life histories, having sessile adults and relying on the planktonic larvae for dispersal. Larvae swim and disperse in a complex fluid environment and the effect of ambient flow on larval behavior could in turn impact their survival and transport. However, to date, most studies on larvae–flow interactions have focused on competent larvae near settlement. We examined the importance of flow on early larval stages by studying how local flow and ontogeny influence swimming behavior in pre-competent larval sea urchins, Arbacia punctulata. We exposed larval urchins to grid-stirred turbulence and recorded their behavior at two stages (4- and 6-armed plutei) in three turbulence regimes. Using particle image velocimetry to quantify and subtract local flow, we tested the hypothesis that larvae respond to turbulence by increasing swimming speed, and that the increase varies with ontogeny. Swimming speed increased with turbulence for both 4- and 6-armed larvae, but their responses differed in terms of vertical swimming velocity. 4-Armed larvae swam most strongly upward in the unforced flow regime, while 6-armed larvae swam most strongly upward in weakly forced flow. Increased turbulence intensity also decreased the relative time that larvae spent in their typical upright orientation. 6-Armed larvae were tilted more frequently in turbulence compared with 4-armed larvae. This observation suggests that as larvae increase in size and add pairs of arms, they are more likely to be passively re-oriented by moving water, rather than being stabilized (by mechanisms associated with increased mass), potentially leading to differential transport. The positive relationship between swimming speed and larval orientation angle suggests that there was also an active response to tilting in turbulence. Our results highlight the importance of turbulence to planktonic larvae, not just during settlement but also in earlier stages through morphology–flow interactions.
  • Article
    Isolating the hydrodynamic triggers of the dive response in eastern oyster larvae
    (John Wiley & Sons, 2015-05-06) Wheeler, Jeanette D. ; Helfrich, Karl R. ; Anderson, Erik J. ; Mullineaux, Lauren S.
    Understanding the behavior of larval invertebrates during planktonic and settlement phases remains an open and intriguing problem in larval ecology. Larvae modify their vertical swimming behavior in response to water column cues to feed, avoid predators, and search for settlement sites. The larval eastern oyster (Crassostrea virginica) can descend in the water column via active downward swimming, sinking, or “diving,” which is a flick and retraction of the ciliated velum to propel a transient downward acceleration. Diving may play an important role in active settlement, as diving larvae move rapidly downward in the water column and may regulate their proximity to suitable settlement sites. Alternatively, it may function as a predator-avoidance escape mechanism. We examined potential hydrodynamic triggers to this behavior by observing larval oysters in a grid-stirred turbulence tank. Larval swimming was recorded for two turbulence intensities and flow properties around each larva were measured using particle image velocimetry. The statistics of flow properties likely to be sensed by larvae (fluid acceleration, deformation, vorticity, and angular acceleration) were compared between diving and non-diving larvae. Our analyses showed that diving larvae experienced high average flow accelerations in short time intervals (approximately 1–2 s) prior to dive onset, while accelerations experienced by non-diving larvae were significantly lower. Further, the probability that larvae dove increased with the fluid acceleration they experienced. These results indicate that oyster larvae actively respond to hydrodynamic signals in the local flow field, which has ecological implications for settlement and predator avoidance.
  • Article
    Departures from isotropy: the kinematics of a larval snail in response to food
    (The Company of Biologists, 2021-01-15) DiBenedetto, Michelle H. ; Meyer-Kaiser, Kirstin S. ; Torjman, Brooke ; Wheeler, Jeanette D. ; Mullineaux, Lauren S.
    The swimming behavior of invertebrate larvae can affect their dispersal, survival and settlement in the ocean. Modeling this behavior accurately poses unique challenges as behavior is controlled by both physiology and environmental cues. Some larvae use cilia to both swim and create feeding currents, resulting in potential trade-offs between the two functions. Food availability is naturally patchy and often occurs in shallow horizontal layers in the ocean. Also, larval swimming motions generally differ in the horizontal and vertical directions. In order to investigate behavioral response to food by ciliated larvae, we measured their behavioral anisotropy by quantifying deviations from a model based on isotropic diffusion. We hypothesized that larvae would increase horizontal swimming and decrease vertical swimming after encountering food, which could lead to aggregation at food layers. We considered Crepidula fornicata larvae, which are specifically of interest as they exhibit unsteady and variable swimming behaviors that are difficult to categorize. We tracked the larvae in still water with and without food, with a portion of the larvae starved beforehand. On average, larvae in the presence of food were observed higher in the water column, with higher swimming speeds and higher horizontal swimming velocities when compared with larvae without food. Starved larvae also exhibited higher vertical velocities in food, suggesting no aggregation behavior. Although most treatments showed strong anisotropy in larval behavior, we found that starved larvae without food exhibited approximately isotropic kinematics, indicating that behavioral anisotropy can vary with environmental history and conditions to enhance foraging success or mitigate food-poor environments.
  • Preprint
    Light stimulates swimming behavior of larval eastern oysters Crassostrea virginica in turbulent flow
    ( 2017-05) Wheeler, Jeanette D. ; Luo, Elaine ; Helfrich, Karl R. ; Anderson, Erik J. ; Starczak, Victoria R. ; Mullineaux, Lauren S.
    Planktonic larvae of the eastern oyster Crassostrea virginica are able to regulate their vertical position in the water, but the environmental cues responsible for this regulation, particularly in turbulent settings, remain unclear. We quantified swimming responses of late-stage oyster larvae in a grid-stirred turbulence tank to determine how light affects the swimming behavior of larvae over a range of hydrodynamic conditions similar to their natural coastal environments. We used particle image velocimetry and larval tracking to isolate larval swimming from local flow and to quantify 3 behavioral metrics: vertical swimming direction, proportion of larvae diving, and proportion of larvae swimming helically. We compared these metrics across turbulence levels ranging from still water (ε = 0 cm2 s-3) to estuarine-like conditions (ε = 0.4 cm2 s-3) in light and dark. At all turbulence levels, light had no effect on the proportion of upward swimming larvae, but elicited detectable increases in the proportion of helical swimming and diving behaviors. We further examined the effect of light and turbulence on specific characteristics of helical trajectories, and found that these environmental cues induce changes to both vertical and horizontal velocities of helically swimming larvae, changing the helix geometry. The increased prevalence of these behaviors in light likely plays an ecological role: increased diving in light (in conjunction with turbulence) is a potential mechanism to enhance settlement success, while changes to helical swimming in light may serve an anti-predatory function. Together, these behaviors provide insight into potentially complex larval responses to multiple simultaneous environmental cues.
  • Article
    Upward swimming of competent oyster larvae Crassostrea virginica persists in highly turbulent flow as detected by PIV flow subtraction
    (Inter-Research, 2013-08-15) Wheeler, Jeanette D. ; Helfrich, Karl R. ; Anderson, Erik J. ; McGann, B. ; Staats, P. ; Wargula, Anna E. ; Wilt, K. ; Mullineaux, Lauren S.
    Investigating settlement responses in the transitory period between planktonic and benthic stages of invertebrates helps shape our understanding of larval dispersal and supply, as well as early adult survival. Turbulence is a physical cue that has been shown to induce sinking and potentially settlement responses in mollusc larvae. In this study, we determined the effect of turbulence on vertical swimming velocity and diving responses in competent eastern oyster larvae Crassostrea virginica. We quantified the behavioural responses of larvae in a moving flow field by measuring and analyzing larval velocities in a relative framework (where local flow is subtracted away, isolating the behavioural component) in contrast to the more common absolute framework (in which behaviour and advection by the flow are conflated). We achieved this separation by simultaneously and separately tracking individuals and measuring the flow field around them using particle image velocimetry in a grid-stirred turbulence tank. Contrary to our expectations, larvae swam upward even in highly turbulent flow, and the dive response became less frequent. These observations suggest that oyster larvae are stronger swimmers than previously expected and provide evidence that turbulence alone may not always be a sufficient cue for settlement out of the water column. Furthermore, at a population level, absolute velocity distributions differed significantly from isolated larval swimming velocities, a result that held over increasing turbulence levels. The absolute velocity distributions indicated a strong downward swimming or sinking response at high turbulence levels, but this observation was in fact due to downwelling mean flows in the tank within the imaging area. Our results suggest that reliable characterization of larval behaviour in turbulent conditions requires the subtraction of local flow at an individual level, imposing the technical constraint of simultaneous flow and behavioural observations.