Anthopleura Xanthogrammica Behavior Studied Utilizing Time-Lapse Photography
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Anthopleura xanthogrammica Behavior Studied Utilizing Time-Lapse Photography By Shannon Reiser A Thesis Presented to the Department of Biology and the Robert D. Clark Honors College in partial fulfillment of the requirements for the degree of Bachelor of Science. November 2013 An Abstract of the Thesis of Shannon Reiser for the degree of Bachelor of Science in the department of Biology to be taken Fall 2013 Title: ANTHOPLEURA XANTHOGRAMMICA BEHAVIOR STUDIED UTILIZING TIME-LAPSE PHOTOGRAPHY Approved: ~Z Date: I~ Aee. 2-0f'r Craigyotltlg epartment of Biology Animals living in a habitat affected by both tidal shifts and day night cycles display a wide variety of behaviors influenced by environmental factors and internal mechanisms. Sessile intertidal invertebrates exhibit extremely slow or subtle behaviors not noticeable during casual observation. This study aimed to observe, analyze, and describe the behaviors associated with the tidal and circadian rhythms of Anthopleura xanthogrammica, giant green anemones, in a tide pool at South Cove, Cape Arago State Park, Oregon. Time lapse video captured using a GoPro camera at a low- to mid-tidal range were used to test the hypothesis A. xamlzogrammica opens on incoming tides. Percent open data were collected from videos and these data were used to evaluate the percentage of animals open in different light conditions. I also examined the data for individual sea anemones to determine if there were individual tendencies. The data suggest that there is a correlation between height of tide and anemone openness. As the tide rises fewer anemones are closed. Additionally the data suggest that anemones are more likely to be closed in direct sunlight. Furthermore, there is a slight, but insignificant difference in average time spent 100% open for each anemone. The most dramatic shifts in behavior occurred in the presence of direct sunlight. ii insignificant difference in average time spent 100% open for each anemone. The most dramatic shifts in behavior occurred in the presence of direct sunlight. Acknowledgements To the multitude of individuals who supported me throughout this entire process, I extend my sincerest gratitude. Professor Young, Professor Sutherland, and Professor Shanks for lending an ear, editing drafts, and keeping me in line. Spring 2014 OIMB students for assisting me in the field. Kate Liddle Broberg, Kerry Mcculloch, Kelsey Ward, Matt Pecenco, Nam Siu, Dominic Pappas, Lauren Gedlinske, and, Maxwell Smith for going above and beyond the call of duty in various ways throughout thesis creation. iii Table of Contents Background _________________________________________ Error! Bookmark not defined. Materials and Methods _________________________________________________ 5 Site Description _____________________________________________________ 5 Field Data Collection _________________________________________________ 7 Analysis ___________________________________________________ 8 Results ____________________________________________________________ 10 Discussion/Conclustion _______________________________ Error! Bookmark not defined.7 Appendix __________________________________________ Error! Bookmark not defined.0 Works Cited ________________________________________ Error! Bookmark not defined.2 iv Background Numerous animal behaviors are dictated by environmentally or internally programmed clocks. Circadian rhythms, or behaviors associated with a 24-hour clock, and circannual rhythms, such as yearly hibernation, are well studied in numerous taxa from unicellular eukaryotes to humans (Tessmar-Raible et al. 2011). But the daily rhythms of marine animals can be based on a more complicated system than that of terrestrial organisms. Some intertidal animals have internal clocks that respond to the tidal cycle as well as the day-night cycle (Palmer 2001). These rhythms are often entrained (initiated) by external (exogenous) cues and maintained (kept ‘on time’) by internal (endogenous) cues. For example, organisms behaving in direct response to changes in pressure or light changes exhibit a clock timed by exogenous cues. On the other hand, endogenous clocks depend on some kind of internal or instinctual mechanism. Some marine organisms behave according to a lunar cycle referred to as a “circatidal” clock (or circalunidian since the tides are caused primarily by the moon). On the west coast tides are semidiurnal with two high and two low tides per “day”(Tessmar-Raible et al. 2011). A tidal day refers to the 24.8-hour interval between moonrises. Two existing schools of thought exist regarding the “mix” of clocks that dictate marine organismal behavior and these hypotheses are at odds with one another. The “circalunidian clock hypothesis” suggests that “two loosely coupled lunidian cycles run in antiphase to produce 12.4 hour intervals” (Wilcockson et al. 2008). This is based on observations of intertidal crab species that experience two activity spikes per day corresponding with the low tides. However, under constant conditions in the lab, the activity peaks disappear after a few days, then reappear, suggesting that the rhythm is not the work of a single circa-tidal oscillator. The other argument postulates a “dedicated 12.4 hour circa-tidal clock” as well as a day/night circadian rhythm. This hypothesis explains the observations that many organisms display behaviors based on both circadian and circatidal schedules. For example, some intertidal invertebrates only spawn during a certain time of the tidal cycle (circatidal), but at night (circadian) (Tessmar-Raible, et al. 2011, Palmer 2001, Wilcockson et al. 2008). Organisms in the intertidal must also grapple with abiotic changes as the tide shifts. A large body of literature shows that temperature changes throughout the tidal cycle can cause severe physiological damage to an organism (Helmuth et al. 2001, Hofmann et al.1995). Intertidal organisms experience changes in temperature, salinity, pressure, turbulence, and food availability each time the tide goes out or comes in (Wilcockson et al. 2008). Therefore organisms have evolved these tidal and daily clocks to improve their chances of surviving rapid environmental changes. For example, many organisms change their physical location when the tide goes out to cope with temperature changes or to avoid desiccation. Other organisms may stay in the same place but employ a physiological protection such as a shell that closes tightly to survive during low tide. Endogenous clocks are well studied in intertidal crab species. For example, it has been shown showed that a fiddler crab, Uca pugnax, maintained the same activity (foraging for food at low tide, immobile at high tide) even in the laboratory under constant light or constant dark (Palmer 2001, Thurman 2004). Furthermore, the crabs shifted this schedule forward 50 minutes every day to correspond with the daily shift in 2 the low and high tides. Field observations reveal that many organisms including, mollusks, arthropods, and fishes display some kind of intertidal migration. An intertidal migration can be as simple as moving from an area of a tide pool exposed to predators and sunlight to an area under cover of nearby rock or algae. An intertidal migration can also cover a relatively large distance. For example, the sea star Pisaster ochraceous travels to a higher zone of the intertidal to prey on Mytilus califonianus, California blue mussels, during high tide, but then must traverse back down to lower tidal heights so as not to be exposed for all of low tide. The distance between mussel beds and water at low tide can sometimes be several meters. Migrating during the day may have several functions including feeding, avoidance of predators, reduction of dessication, and reproduction. Feeding in some animals is only possible when the intertidal zone is submerged or when the tide has receded. Moreover, migration in some species avoiding predators only occurs when high or low tide is at night (Gibson 2003). Recent research revealed that another anemone species, Nematostella vectensis, maintains a 24 hour behavioral cycle in which locomotion is predominantly nocturnal. Under constant light or constant dark conditions, the timing of locomotory behavior remained the same as in day/night conditions, suggesting the presence of some sort of endogenous mechanism (Hendricks et al 2012). Anthopleura xanthogrammica, the study organism for the present research is a sessile invertebrate from phylum Cnidaria. Anthopleura xanthogrammica inhabits mid to lower tidal regions in cracks, crevices, in pools, and on rocks and is distinguishable by its size, bright green color, and ring of tentacles around its oral disc. A. xanthogrammica feeds on invertebrates, particularly mussels, dislodged by waves or by 3 the foraging activity of P. ochraceous. This allows me to postulate that anemones will be open more frequently when the tide is higher (Sebens 1986). The gastrodermal tissues of A. xanthogrammica contain at least one type of symbiotic algae that require light for photosynthesis. This leads to the prediction that anemones respond to both changes in tide (as they are more likely to catch prey at high tide), and to light because sunnier conditions provide an opportunity to expose symbionts to sunlight (Trench, 1971, Muscantine et al 1975). Research on a congeneric anemone Anthopleura elegantissma, the aggregating anemone, revealed that presence or absence of symbiotic zooxanthellae affected their behavior. Those with the symbiont clustered near a light in an