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Magnetic Sense of Juvenile Gulf Flounder

(

Paralichthys albigutta

)

By

EmmaLi G. Tsai

Senior Honors Thesis

Environment, Ecology, and Energy Program

University of North Carolina at Chapel Hill

(26 November 2019)

Approved:

Dr. Joel Fodrie, Thesis Advisor

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ABSTRACT... 3

1. INTRODUCTION... 4

2. METHODOLOGY... 7

2.1 Fish Capture and Husbandry... 7

2.2 Magnetic Coil Structure... 8

2.3 Experimental Conditions... 9

2.4 Experimental Treatments... 10

2.4.1 Magnetic Compass Experiments... 10

2.4.2 Magnetic Map Experiments... 11

2.5 Experimental Procedure... 13

2.6 Data Analysis... 13

3. RESULTS... 15

3.1 Magnetic Compass Results... 15

3.2 Magnetic Map Results... 16

4. DISCUSSION... 17

4.1 Magnetic Compass Experiments... 17

4.2 Magnetic Map Experiments... 18

5. FUTURE STEPS... 19

6. APPLICATIONS & IMPLICATIONS... 21

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ABSTRACT

Diverse marine taxa use Earth's magnetic field for navigation. This ability, known as

magnetic navigation, has classically been grouped into two main types – the magnetic “compass”

and “map” sense. Organisms possessing a magnetic compass sense are able to use the Earth’s

magnetic field to gain directional information, similar to a human compass. In contrast, an

organism possessing a magnetic map sense uses the Earth’s magnetic field to determine their

position relative to a goal, similar to a human GPS. We assessed for the presence of both a

magnetic compass and map sense in juvenile Gulf Flounder (Paralichthys albigutta), a species of

migratory flatfish found in coastal waters spanning from North Carolina to the Gulf of Mexico.

In both experiments, a magnetic coil system was used to produce different magnetic field

treatments. To test for a magnetic compass sense, flounder were tested in one of two magnetic

treatments: (1) the ambient magnetic field of the test site; (2) the same magnetic field but with

reversed polarity, where the direction of magnetic north was reversed. To search for the presence

of a magnetic map sense, we tested flounder in one of two magnetic fields that exist at two

locations: (1) north of the test site (near New Jersey); and (2) south of the test site (near

Jamaica). Flounder consistently oriented towards a magnetic direction in our compass

experiments, which suggests the presence of a compass sense. In the magnetic map experiments,

flounder exhibited orientations that were significantly different between northern and southern

treatments, providing evidence for a magnetic map sense. Considering past magnetic navigation

research has mainly focused on long-distance migrants, the discovery of a both a magnetic

compass and map sense in flatfish brings to light how these senses could be useful for organisms

undergoing relatively shorter migrations between nearshore and offshore environments.

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1. INTRODUCTION

Many fish species move between ocean habitats that are critical for different stages of

their life history (Aas et al., 2011; Dingle, 2014). In fish, an abundance of these history strategies

involve offshore larval dispersal, a pelagic stage, and subsequent settlement in a coastal nursery

habitat where fish continue their development (Dingle, 2014). As knowledge behind fish

migration has developed, past biotelemetry and stable isotope analysis has discovered these

movements can actually cover significant distances across ocean environments (Bultel et al.,

2014; Madigan et al., 2014). During these long-distance migrations, fish utilize a suite of

navigational systems to successfully arrive at habitats essential for growth and reproduction

(Gould, 1998). The systems must be reliable to account for directional errors that inevitably arise

when navigating an ever-changing ocean environment (Lohmann, 2008). However, obtaining

consistent navigational information is a challenge in the open-ocean environment; a habitat

largely devoid of visual landmarks. How fish are able to achieve these long-distance migrations

can be elucidated by understanding the sensory mechanisms guiding these movements.

Previous research suggests that fish use an amalgamation of acoustic, visual,

hydrodynamic, olfactory, and geomagnetic cues to navigate the ocean environment (Radford et

al., 2010; Gardiner & Atema, 2014; Groves et al., 1968; Lohmann 2008). In such an

environment, information gained from Earth’s magnetic field can be particularly beneficial to

migrating organisms. It is present everywhere on the planet – from the deepest ocean trenches to

the highest mountains, and largely undisturbed by extreme weather events (Lohmann et al.,

2004). Using the Earth’s magnetic field for navigational information is known as “magnetic

navigation”, and is an ability that has been discovered in diverse taxa ranging from birds,

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Vácha et al., 2009; Kimchi & Terkel, 2001; Ernst & Lohmann, 2018; Naisbett-Jones et al., 2017;

Putnam et al., 2014; Lohmann et al., 2004). As knowledge behind this ability has developed,

there are two main types of magnetic sense that have been identified by scientists. The first is

known as the “magnetic compass” sense, where an organism is able to use the Earth’s magnetic

field to identify a specific magnetic direction to maintain a constant heading, similar to a human

compass (Lohmann et al., 1993). This ability has been identified in many organisms, such as

lobsters, fish, and mollusks (Ernst & Lohmann, 2018; Bottesch et al., 2016; Cain et al., 2005).

The second is known as a “magnetic map” sense, where an organism can use the Earth’s

magnetic field to determine their position relative to a goal, similar to a human GPS (Wiltschko

& Wiltschko, 2005). The magnetic map sense has been discovered in many long-distance

migrants, such as eels, sea turtles, and salmon (Naisbett-Jones et al., 2017; Lohmann et al., 2004;

Putnam et al., 2017). While our understanding of magnetic navigation has increased substantially

over the years, current research has largely focused on iconic long-distance migrants that exhibit

high fidelity to specific locations. Comparatively little research, however, has been conducted on

fish possessing other life history characteristics.

Fish that do not exhibit high fidelity to specific locations

nonetheless experience navigational obstacles where a magnetic sense

could provide useful information. An example of such a case would be

flatfish (Pleuronectiformes spp.); an order of oceanadromous demersal

fish that face the challenge of navigating the ocean floor. Flatfish

species adopt a relatively common life history strategy that requires

migration of 10s-100s km between inshore and offshore habitats (Gibson et al., 2015). Adult

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producing small larvae. These pelagic larvae develop in the water column and undergo an

incredible metamorphosis to become demersal juveniles, where one eye fully migrates to the

other side of the head (Figure 1; McMenamin & Parichy, 2013). During the late spring and early

summer months, juveniles settle in continental nursery habitats to continue their development.

During the offseason, flatfish either migrate offshore or southward to reach warmer waters

(Gibson et al., 2015). While previous telemetry studies with European plaice (Pleuronectes

platessa) have demonstrated that this species of flatfish are able to maintain a constant compass

bearing, to our knowledge no study has researched the magnetic sense in any species of flatfish

(Metcalfe et al., 1993).

Here, we investigated for the presence of both a magnetic compass and map sense in

juvenile Gulf Flounder (Paralichthys albigutta). Gulf flounder is a species of flatfish commonly

found in coastal waters across the southeastern United States, ranging from North Carolina to an

undetermined southern limit around the Gulf of Mexico (Figure 2). For the magnetic compass

experiments that I assisted with, we used two separate treatments: (1) the ambient magnetic field

present at Morehead City, NC; and (2) the ambient magnetic field but with reversed polarity. In

order to assess the presence of a magnetic map sense, I modified magnetic field parameters to

represent magnetic fields at two geographic locations: (1) a magnetic field found north near New

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2. METHODOLOGY

2.1 Fish Capture and Husbandry

Gulf Flounder ranging from 8-32 cm were obtained via trawling and seining seagrass

beds in Back Sound, North Carolina (Figure 3; 34.68° N, 76.56° W) during July 3rd-10th of

2018 for the compass experiments, and May 24th - June 6th of 2019 for both the compass and

map experiments. After capture in the field, fish were kept in a non-magnetic aerated container

that was replenished with local water every hour. After transportation, fish were maintained in

elevated non-magnetic outdoor tanks located at the University of North Carolina’s Institute of

Marine Science (UNC-IMS; 34.72° N, 76.75° W). These tanks were elevated to reduce the

magnetic influence of potentially buried objects, and isolated to reduce human-generated

magnetic anomalies. During this time in captivity, the fish were fed small anchovies every other

day and supplied with local water from Bogue Sound that mimicked natural conditions. In the

absence of adverse weather, collected flounder were tested between 24-48 hours after capture to

avoid stress associated with prolonged time spent in captivity. Flounder collected from these

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Figure 3. Map showing collection site for Gulf Flounder near Morehead City, North Carolina. Dark blue dotted oval displays the sea grass collection sites located in Bogue Sound, North Carolina. The red star indicates the location

where all experiments took place, UNC-IMS.

2.2 Magnetic Coil Structure

In order to produce the magnetic treatments used in both the compass and map

experiments, we utilized a Merritt et al. (1983) 4-Coil design, which is a wooden structure that

supports 6,000 feet of vertically and horizontally wrapped insulated copper wire (Figure 4).

These copper wires were connected to a BK Precision Model 1550 DC Power Supply box and

supplied with constant current in order to produce a constant magnetic field throughout the

duration of our experiments. By running a current through these vertically and horizontally

wrapped wires, the magnetic coil structure can be used to manipulate the ambient magnetic field

present at the UNC-IMS. All of our treatments were based on magnetic parameters produced by

NOAA’s International Geomagnetic Reference Field (IGRF-12) and verified using a FVM400

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All fish were tested on the experimental platform located inside of the wooden structure

and within enclosed black-out curtains. Black-curtains were used to remove all celestial cues and

isolate the magnetic response from the flounder. This experimental platform was also centered

inside of the structure where the magnetic field produced was uniform, and lifted to avoid

magnetic disturbance from objects or debris that may be buried under the coil structure.

2.3 Experimental Conditions

Our preliminary studies with Gulf Flounder illustrated higher activity levels at night,

which is consistent with previous experiments on similar flatfish species (Miyazaki et al., 1997).

Therefore, all experiments were conducted at night between 20:50-4:20. Four opaque circular

orientation arenas were placed on the raised magnetic coil platform and enclosed with black-out

curtains (Figure 5). These arenas were non-magnetic and isolated from each other to reduce

disruption resulting from adjacent flounder movements. Each orientation arena was filled at

uniform depth with local water obtained from Bogue Sound, and large black arrows were drawn

on the coil platform to reference magnetic north. In order to film flounder behavior, an infrared

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Figure 5. Image of the experimental platform where all experiments were filmed. Each of the four orientation arenas are isolated and filled to a certain depth with water obtained

from Bogue Sound. The black arrows on the platform are directed towards magnetic north

and were used for video analysis.

2.4 Experimental Treatments

2.4.1 Magnetic Compass Experiments

All compass experiments were conducted between July 3rd-10th of 2018 and May 24th

-June 6th of 2019 outside of UNC-IMS. Flounder were tested in two experimental treatments: (1)

the ambient magnetic field located at Morehead City, NC, and (2) a magnetic field with reversed

polarity where magnetic north would be present on the opposite end of the experimental

platform. The act of reversing magnetic north is often used as a method for determining the

presence of a compass sense in previous experiments (Lohmann & Lohmann, 1993; Lohmann et

al., 1995). If flounder possess a magnetic compass sense, we would hypothesize that flounder

experiencing the reversed magnetic north treatment would orient in approximately the opposite

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aimed to gather a total sample of roughly 20 flounder for each treatment, which is consistent

with previous magnetic compass experiments (Cresci et al., 2017; Durif et al., 2013).

(a) (b)

Figure 6. Our expected results if Gulf Flounder possess a magnetic compass sense. (a) In the ambient Morehead City field, the flounder orient towards a

specific magnetic direction, such as west. (b) Upon reversal of the field, the

flounder should orient towards approximately the opposite direction, but still

towards the same magnetic direction.

2.4.2 Magnetic Map Experiments

All magnetic map experiments occurred between the June 23rd - August 13th of 2019. For

the magnetic map experiments, I modified both magnetic field intensity and the angle the

magnetic field intersects the surface of the Earth (inclination), to represent magnetic fields of

different geographic locations. I generated magnetic fields that would be found at locations

approximately 600km north and south of our experimental location of North Carolina: (1) a

magnetic field of stronger field intensity and larger inclination angle found north near New

Jersey, and (2) a magnetic field of weaker field intensity and smaller inclination angle found

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tested organisms in magnetic fields present at different geographic locations (Naisbett-Jones et

al., 2017; Boles & Lohmann, 2003; Putman et al., 2014). If flounder possess a magnetic map

sense, I hypothesize that flounder would respond to changes in magnetic field intensity and

inclination and exhibit significantly different orientations between experimental treatments. It is

possible they would orient in approximately opposite directions in order to return to the magnetic

field found near their caught location of North Carolina (Figure 7). I aimed to gather a sample

size of approximately 50 fish per treatment, which is consistent with previous magnetic map

experiments (Fischer et al., 1999).

Figure 7. My expected results if Gulf Flounder possess a magnetic map sense. The northern New Jersey treatment has a magnetic field intensity of approximately 51.1 µT and inclination of 65°, and the southern Jamaica treatment

has a magnetic field intensity of approximately 38.4 µT and inclination of 46°. Flounder should sense these changes

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treatments. The fish could orient where they would hypothetically return to the magnetic field intensity of 48.6 µT

and inclination of 61° present at their caught location of Morehead City, NC.

2.5 Experimental Procedure

The experimental procedure was similar for both the magnetic compass and magnetic

map experiments. In both experiments, treatments were interspersed throughout the night to

control for temporal variation, such as tidal cycle, which has been shown to previously influence

magnetic orientation in other aquatic species (Cresci et al., 2017). Prior to the start of an

experiment, the four orientation arenas were filled at uniform depth with local water from Bogue

Sound and the particular magnetic treatment was generated. Fish were transported from their

outdoor tank in a non-magnetic bucket to the coil structure, carefully placed into an orientation

arena, and given a 10 - minute acclimation period. Flounder behavior was filmed for a total

period of 2 hours for the compass experiments, and 1.5 hours for the map experiments. The

magnetic compass experiments were filmed for a longer duration than the map experiments to be

consistent with experiments conducted in 2018, which used generally larger and more sedentary

flounder. Between experiments, the orientation arenas were emptied and replenished with local

water, the coil was configured to generate the magnetic field for the other treatment, and four

new fish were placed into each orientation arena. Each fish was only tested once in response to

an experimental treatment and a single individual was either used for the magnetic compass or

map experiments, but not both. After testing, flounder were released into Bogue Sound.

2.6 Data Analysis

We used flounder “oriented swimming movement” as the behavioral assay to determine

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that caused the flounder to swim vertically or come in contact with the perimeter of the

orientation arena for an extended period of time. Fish that did not move for the duration of the

experiment were not included in the analysis. Following the acclimation period, the beginning of

the first 10 directional movements made by each fish were documented, and the angle of this

movement relative to magnetic north was measured using ImageJ software. Any movements

made after the first 10 were determined to be more exploratory rather than directional, and

therefore excluded from the analysis. The angles of movement for a single flounder were

averaged to compute a mean angle of directional preference for a single fish throughout the

duration of the experiment, and graphed using the circular statistics software, Oriana. Using

Oriana, it was possible to conduct a Rayleigh test to search for significant unimodal orientation

within experimental group, and a Mardia-Watson-Wheeler test to assess significance between

experimental groups (Wilkie 1983; Batschelet 1981). Across all tests, a P-value less than 0.05

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3. RESULTS

3.1 Magnetic Compass Results

Figure 8. Each dot represents the mean angle of movement for a single flounder and the dashed black lines display 95% confidence intervals. (a) In the Earth’s magnetic field treatment, flounder significantly oriented west (N=20;

Mean Angle= 282°; Rayleigh P= 2.09E-4). (b) In the reversed magnetic north field, flounder significantly oriented

in the approximately the opposite direction, but towards the same magnetic direction (N=23; Mean Angle=72.2°;

Rayleigh P= 5.70E-4).

In the ambient magnetic field present at the experimental location, flounder significantly

oriented towards magnetic west (Figure 8a). Upon reversal of the field, flounder significantly

oriented in approximately the opposite direction, but still towards the same magnetic direction

(Figure 8b). The orientation responses of these two treatments were significantly different from

each other (Mardia-Watson-Wheeler P= 5.63E-5). In both treatments, flounder orientation data

gathered from 2018 and 2019 were not significantly different from one another, and tidal state

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3.2 Magnetic Map Results

Figure 9. Each dot represents the mean angle of movement for a single fish, and 95% confidence intervals are displayed as dashed black lines. The northern New Jersey magnetic field treatment is illustrated above in red, and

the southern Jamaica magnetic field treatment in blue. Average direction of movement for each treatment is

indicated by a black arrow. (a) In the northern New Jersey treatment, fish significantly oriented towards magnetic

southeast (N=48; Mean Angle= 147°; Rayleigh P= 0.022). (b) In the southern Jamaica treatment, fish exhibited

orientations that could not be distinguished from random (N=51; Rayleigh P=0.148).

In the northern New Jersey treatment with a stronger magnetic field intensity and larger

inclination angle, flounder significantly oriented towards magnetic southeast (Figure 9a). In the

Jamaica treatment with a weaker magnetic field intensity and smaller inclination angle, fish

exhibited orientations that could not be distinguished from random (Figure 9b). Orientations

between these two experimental treatments were significantly different from one another

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4. DISCUSSION

Our results suggest that Gulf Flounder possess the ability to use Earth’s magnetic field

for both directional and positional information. In the magnetic compass experiments, fish

oriented towards magnetic west in both the ambient and reversed magnetic field treatments.

Maintaining consistent magnetic orientation even upon reversal of the field illustrates the

presence of a magnetic compass sense, where an organism can use the magnetic field for

directional information. In my magnetic map experiments, flounder exhibited significantly

different orientations between the northern and southern magnetic map treatments. Considering

flounder responded differently to changes in field intensity and inclination angle, these results

suggest that Gulf Flounder possess the ability to determine positional information from Earth’s

magnetic field, or a magnetic map sense.

4.1 Magnetic Compass Experiments

The results of our magnetic compass experiments with Gulf Flounder were consistent

with previous research, where an organism possessing a compass sense was able to consistently

orient towards the same magnetic direction (Lohmann & Lohmann, 1993; Lohmann et al., 1995).

This sense has been discovered in diverse marine organisms, such as reef fish and isopods, and

may provide sufficient directional information to guide organisms between habitats (Bottesch et

al., 2016; Ugolini & Pezzani, 1995). Building off of existing literature, our discovery of a

compass sense in Gulf Flounder illuminates how this sense could also be useful to migratory

flatfish that travel between nearshore and offshore environments. In particular, our results could

explain the consistent directional heading of European plaice, a genetically similar flatfish

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There are a couple of possible interpretations of the western directional preference

flounder exhibited in the magnetic compass experiments. First, it is possible that this westerly

preference may be a recruitment response. Considering we collected these flounder during the

summer when they would normally be recruiting, orienting westward may be sufficient to guide

flounder to coastal nursery habitats. However, flounder may also be responding to this cue on a

more local scale. It is possible they may be orienting westward in order to travel along Bogue

Sound, a body of water located near the experimental location of UNC-IMS. Future experiments

should be conducted to determine the scale that flounder are responding to magnetic information.

4.2 Magnetic Map Experiments

In the magnetic map experiments, I found that fish experiencing the northern New Jersey

magnetic field significantly oriented towards magnetic southeast, a direction that would

hypothetically allow them to return to their caught location of Morehead City, NC. Flounder

experiencing the southern magnetic field treatment near Jamaica exhibited orientations that could

not be distinguished from random. I believe these different responses could be explained by the

known distribution of Gulf Flounder, which extends from North Carolina to an undetermined

southern limit near the Gulf of Mexico. Since the New Jersey magnetic field treatment extends

far north above their natural range, flounder would recognize this magnetic field as present in an

uninhabitable environment. The New Jersey coastline is colder than the locations Gulf Flounder

usually occupy, and it would come at a high cost for an individual to dwell in this location.

Therefore, flounder exhibited a strong southern response to leave this magnetic field and return

to a location they could inhabit. This is consistent with previous map experiments, where testing

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elicit a strong directional response (Putnam et al., 2014). Additionally, given there is anecdotal

evidence of Gulf Flounder near Jamaica, our southern magnetic field is present in a location that

flounder could feasibly occupy. It is possible that our southern Jamaica field may not have been

outside of the southernmost limit of Gulf Flounder’s range to elicit a northern response, leading

to random orientation.

5. FUTURE STEPS

Previous magnetic navigation research has largely focused on discrete life history stages

of an organism. However, to date, no research has ever been conducted that offered a more

comprehensive view of how the use of this sense transforms throughout ontogeny. Considering

fish at varying have history stages have different objectives, the magnetic sense is likely used

differently throughout an individual’s lifetime. For flounder, it would be fascinating to illustrate

how this sense could be recapitulated in younger life history stages, especially in light of new

empirical evidence of magnetic navigation in Atlantic Haddock larvae and reef fish larvae

(Cresci et al., 2019; O'Connor & Muheim, 2017; Bottesch et al., 2016). Searching for the

presence of a magnetic sense in flounder larvae could illustrate how they may have a more active

role in controlling the trajectory of their recruitment, thereby enhancing in our understanding of

larval dispersal and population connectivity. In the adults, investigating the transformation of the

magnetic sense could elucidate how this sense may be used when navigating offshore towards

spawning locations. Ultimately, understanding the use of the magnetic sense throughout

ontogeny could help us better understand the senses guiding flatfish movements during different

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While examining different stages of life history would illustrate the transformation of the

magnetic sense through an individual’s lifetime, another fascinating avenue of research would be

to assess the influence of seasonality on directional preference. Seasonal shifts in magnetic

directional preference have previously been discovered in migratory bird species, such as the

European robin, and migrating moths (Wiltschko & Wiltschko, 2001; Chapman et al., 2008).

This has also demonstrated with aquatic migratory species, where temperature was discovered to

be a large determinant in the strength and directional preference of a species of eel (Durif et al.,

2013). Considering Gulf Flounder either migrate offshore or southwards during the offseason, it

is possible that seasonality and thermal change could transform their summer western response

into a southern response during the fall and winter months. Future research should be conducted

to further investigate the possibility of this seasonal behavior in Gulf Flounder.

Aside from season, the tidal cycle is another possible parameter that has been shown to

influence the directional preference in magnetoreceptive organisms. Many aquatic organisms

utilize tidal change as a method of transport between near and offshore environments

(Tankersley et al., 1998; Gibson, 2003). Recent empirical research that suggests that this method

of transport may be linked to the magnetic compass sense (Cresci et al., 2017). Given tidal

transport has also been discovered in previous tracking studies with European plaice, a

genetically similar flatfish species, it is possible that Gulf Flounder may possess a similar system

(Metcalfe et al., 1990). Understanding how the magnetic compass sense is influenced by tidal

change would provide insight on how flounder may use the tidal cycle to conserve energy and be

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6. APPLICATIONS & IMPLICATIONS

Past research on magnetic navigation has largely focused on organisms that undergo

long-distance migrations along complex migratory routes. Discovering a magnetic sense in

flatfish demonstrates how this sense could provide useful navigational information for organisms

that undergo shorter migrations between nearshore and offshore environments. Moreover,

flatfish exhibit a rather common life history strategy comprised of larval dispersal and

subsequent recruitment to coastal habitats. Finding a magnetic sense in flatfish illustrates how

this sense may be present in other organisms that possess similar life history strategies. On a

broader scale, this information serves as a small puzzle piece to the larger developing field of

magnetic sensory research in attempt to deepen the understanding of how sensory cues are

involved in long-distance migrations.

As magnetic sensory research has developed, the processes behind how organisms are

able to detect the Earth’s magnetic field is still a question that still remains unanswered by

scientists. Since the magnetic field passes through the whole organism, the processes behind this

sense could theoretically be present anywhere in biological tissue (Johnsen & Lohmann, 2005).

A couple of hypotheses have been developed in an effort to explain this phenomenon from

observing a number of different organisms. Scientists have found evidence that birds are able to

determine their geographic position by using radical pair-forming molecules in their eyes that

communicate with cryptochrome contained in their beaks (Wiltschko & Wiltschko, 2005).

Elasmobranchs, such as rays and sharks, detect magnetic variations by using an induction-based

method that involves monitoring fluctuations in current generated through their electric organs

(Akoev, 1990). Bats have also been hypothesized to be able to use magnetite and a series of

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Flounder’s transparent larval stage provides an interesting opportunity for scientists to search for

these processes within a smaller organism. Their transparency is particularly useful because it

allows for easier scanning for magnetic particles, such as magnetite, that may aid in this sensing

ability (Holland et al., 2008). Overall, the lifecycle and physiological characteristics of Gulf

Flounder allow them to serve as an ideal mechanistic model to resolve the enigma of the physical

processes behind magnetoreception.

Besides being a model organism to search for the magnetoreceptor, many species of

flatfish support important commercial fisheries. In North Carolina, both summer and southern

flounder were found by the Division of Marine Fisheries to be two of the top five fished species

in 2017 and their decline prompted stronger fishing regulations (“NC Flounder Limits”, 2018).

Understanding the senses that guide flatfish movements and migrations could allow for the

construction of successful fishery management decisions, thereby investing in the long-term

prosperity of the flatfish stock and the economies that depend on it.

Gulf flounder possessing both a magnetic compass and map sense could help explain

how flounder navigate the environment they occupy, but also highlights the possible influence

that human magnetic equipment can have on magnetoreceptive organisms. Offshore wind energy

transfers large amounts of current through undersea cables back to the coast, which creates

anomalies in the magnetic field surrounding them (Öhman et al., 2007). This is of particular

concern for flounder, who occupy the ocean floor and could be strongly influenced by these

magnetic anomalies during their nearshore and offshore migrations. Aside from flounder, these

disruptions in the magnetic field have an unknown influence on the orientation, behavior, and

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ramifications need to be further explored by scientists and should be taken into consideration

concerning future developments.

REFERENCES

Aas, Ø., Einum, S., Klemetsen, A., Skurdal, J., (2011). Atlantic salmon ecology. Chichester: Wiley-Blackwell.

Akoev, G. N. (1990). Temperature Sensitivity of the Ampullae of Lorenzini of Elasmobranchs. Thermoreception and Temperature Regulation,44-52. doi:10.1007/978-3-642-75076-2_6

Batschelet, E. (1981). Circular statistics in biology. New York: Academic Press.

Boles, L. C., & Lohmann, K. J. (2003). True navigation and magnetic maps in spiny lobsters. Nature, 421(6918), 60–63. doi: 10.1038/nature01226

Bottesch, M., Gerlach, G., Halbach, M., Bally, A., Kingsford, M. J., & Mouritsen, H. (2016). A magnetic compass that might help coral reef fish larvae return to their natal reef. Current Biology, 26(24). doi: 10.1016/j.cub.2016.10.051

Bultel, E., Lasne, E., Acou, A., Guillaudeau, J., Bertier, C., & Feunteun, E. (2014). Migration behaviour of silver eels (Anguilla anguilla) in a large estuary of Western Europe inferred from acoustic telemetry. Estuarine, Coastal and Shelf Science, 137, 23–31. doi: 10.1016/j.ecss.2013.11.023

Cain, S. D., Boles, L. C., Wang, J. H., & Lohmann, K. J. (2005). Magnetic Orientation and Navigation in Marine Turtles, Lobsters, and Molluscs: Concepts and Conundrums. Integrative and Comparative Biology, 45(3), 539-546. doi:10.1093/icb/45.3.539

Chapman, J. W., Reynolds, D. R., Hill, J. K., Sivell, D., Smith, A. D., & Woiwod, I. P. (2008). A seasonal switch in compass orientation in a high-flying migrant moth. Current Biology, 18(19). doi: 10.1016/j.cub.2008.08.014

Cresci, A., Paris, C. B., Durif, C., Shema, S., Bjelland, R. M., Skiftesvik, A. B., & Browman, H. I. (2017). Glass eels (Anguilla anguilla) have a magnetic compass linked to the tidal cycle. Science advances, 3(6), e1602007. doi:10.1126/sciadv.1602007

Gould, J. L. (1998). Sensory bases of navigation. Current Biology, 8(20), R731–R738. https://doi.org/https://doi.org/10.1016/S0960-9822(98)70461-0

Dingle, H. (2014). Migration the biology of life on the move. Oxford: Oxford University Press.

Durif, C. M., Browman, H. I., Phillips, J. B., Skiftesvik, A. B., Vøllestad, L. A., & Stockhausen, H. H. (2013). Magnetic Compass Orientation in the European Eel. PLoS ONE, 8(3). doi:10.1371/journal.pone.0059212

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Fischer, J., Freake, M., Borland, S., & Phillips, J. (2001). Evidence for the use of magnetic map information by an amphibian. Animal Behaviour, 62(1), 1–10. doi: 10.1006/anbe.2000.1722

Gardiner, J. M., & Atema, J. (2014). Flow Sensing in Sharks: Lateral Line Contributions to Navigation and Prey Capture. Flow Sensing in Air and Water, 127–146. doi: 10.1007/978-3-642-41446-6_5

Gibson, R. N. (2003). Go with the flow: tidal migration in marine animals. Migrations and Dispersal of Marine Organisms, 153–161. doi: 10.1007/978-94-017-2276-6_17

Gibson, R. N., Nash, R. D., Geffen, A. J., & W., V. D. (2015). Flatfishes: Biology and exploitation. Chichester, West Sussex, UK: Wiley Blackwell.

Gould, J. L., & Gould, C. G. (2012). Natures compass: The mystery of animal navigation. Princeton: Princeton University Press.

Groves, A. B., Collins, G. B., & Trefethen, P. S. (1968). Roles of Olfaction and Vision in Choice

of Spawning Site by Homing Adult Chinook Salmon (Oncorhynchus tshawytscha). Journal

of the Fisheries Research Board of Canada, 25(5), 867–876. doi: 10.1139/f68-082

Holland, R. A., Kirschvink, J. L., Doak, T. G., & Wikelski, M. (2008). Bats Use Magnetite to Detect the Earths Magnetic Field. PLoS ONE,3(2). doi:10.1371/journal.pone.0001676

Johnsen, S., & Lohmann, K. J. (2005). The physics and neurobiology of magnetoreception. Nature Reviews Neuroscience, 6(9), 703-712. doi:10.1038/nrn1745

Kimchi, T., Terkel, J. (2001). Magnetic compass orientation in the bling mole rat Spalax ehrenbergi. Journal of Experimental Biology, 204, 751-758.

Lohmann, K. J., & Lohmann, C. (1993). A Light-Independent Magnetic Compass in the Leatherback Sea Turtle. The Biological Bulletin, 185(1), 149-151. doi:10.2307/1542138

Lohmann, K. J., Lohmann, C. M., Ehrhart, L. M., Bagley, D. A., & Swing, T. (2004). Animal behaviour: Geomagnetic map used in sea-turtle navigation. Nature,428(6986), 909-910. doi:10.1038/428909a

Lohmann, K. J., Putman, N. F., & Lohmann, C. M. (2008). Geomagnetic imprinting: A unifying hypothesis of long-distance natal homing in salmon and sea turtles. Proceedings of the National Academy of Sciences, 105(49), 19096-19101. doi:10.1073/pnas.0801859105

Lohmann, K. J., Pentcheff, N. D., Nevitt, G. A., Stetten, G. D., Zimmer-Faust, R. K., Jarrard, H. E., & Boles, L. C. (1995). Magnetic Orientation of Spiny Lobsters in the Ocean: Experiments with Undersea Coil Systems. The Journal of Experimental Biology. Retrieved December 12, 2018.

(25)

Mcmenamin, S. K., & Parichy, D. M. (2013). Metamorphosis in Teleosts. Current Topics in Developmental Biology Animal Metamorphosis, 127–165. doi: 10.1016/b978-0-12-385979-2.00005-8

Merritt, R., Purcell, C., & Stroink, G. (1983). Uniform magnetic field produced by three, four, and five square coils. Review of Scientific Instruments, 54(7), 879-882. doi:10.1063/1.1137480

Metcalfe, J. D., Arnold, G. P., & Webb, P. W. (1990). The energetics of migration by selective tidal stream transport: an analysis for plaice tracked in the southern North Sea. Journal of the Marine Biological Association of the United Kingdom, 70(1), 149–162. doi: 10.1017/s0025315400034275

Metcalfe, J. D., Holford, B. H., & Arnold, G. P. (1993). Orientation of plaice (Pleuronectes platessa) in the open sea: Evidence for the use of external directional clues. Marine Biology, 117(4), 559-566. doi:10.1007/bf00349766

Miyazaki, T., Masuda, R., Furuta, S., & Tsukamoto, K. (1997). Laboratory. Observation on the Nocturnal Activity of Hatchery-reared Juvenile Japanese Flounder Paralichthys olivaceus. Fisheries Science, 63(2), 205–210. doi: 10.2331/fishsci.63.205

Naisbett-Jones, L. C., Putman, N. F., Stephenson, J. F., Ladak, S., & Young, K. A. (2017). A Magnetic Map Leads Juvenile European Eels to the Gulf Stream. Current Biology,27(8), 1236-1240. doi:10.1016/j.cub.2017.03.015

NC Flounder Limits Have Changed. (2018, June 12). Retrieved from https://islandlifenc.com/nc-flounder-limits-changed/

O’Connor, J. O., & Muheim, R. (2017). Pre-settlement coral-reef fish larvae respond to magnetic field changes during the day. 2874–2877. https://doi.org/10.1242/jeb.159491

Öhman, M. C., Sigray, P., & Westerberg, H. (2007). Offshore Windmills and the Effects of Electromagnetic Fields on Fish. AMBIO: A Journal of the Human Environment, 36(8), 630-633. doi:10.1579/0044-7447(2007)36[630:owateo]2.0.co;2

Putman, N., Scanlan, M., Billman, E., O’Neil, J., Couture, R., Quinn, T., . . . Noakes, D. (2014). An Inherited Magnetic Map Guides Ocean Navigation in Juvenile Pacific Salmon. Current Biology,24(4), 446-450. doi:10.1016/j.cub.2014.01.017

Radford, C. A., Stanley, J. A., Simpson, S. D., & Jeffs, A. G. (2011). Juvenile coral reef fish use sound to locate habitats. Coral Reefs, 30(2), 295–305. doi: 10.1007/s00338-010-0710-6

Tankersley, R. A., Wieber, M. G., Sigala, M. A., & Kachurak, K. A. (1998). Migratory Behavior of Ovigerous Blue Crabs Callinectes sapidus: Evidence for Selective Tidal-Stream Transport. The Biological Bulletin, 195(2), 168–173. doi: 10.2307/1542824

(26)

Vacha, M., Puzova, T., & Kvicalova, M. (2009). Radio frequency magnetic fields disrupt magnetoreception in American cockroach. Journal of Experimental Biology, 212(21), 3473– 3477. doi: 10.1242/jeb.028670

Wilkie, D. (1983). Rayleigh Test for Randomness of Circular Data. Applied Statistics, 32(3), 311. doi:10.2307/2347954

Wiltschko, W., Gesson, M., & Wiltschko, R. (2001). Magnetic compass orientation of European robins under 565 nm green light. Naturwissenschaften, 88(9), 387-390. doi:10.1007/s001140100248

Figure

Figure 3. Map showing collection site for Gulf Flounder near Morehead City, North Carolina
Figure 5. Image of the experimental platform where all experiments were filmed. Each of
Figure 6. Our expected results if Gulf Flounder possess a magnetic compass
Figure 7. My expected results if Gulf Flounder possess a magnetic map sense. The northern New Jersey treatment
+3

References

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