Magnetic maps in animal navigation.

In addition to providing animals with a source of directional or 'compass' information, Earth's magnetic field also provides a potential source of positional or 'map' information that animals might exploit to assess location. In less than a generation, the idea that animals use Earth's magnetic field as a kind of map has gone from a contentious hypothesis to a well-established tenet of animal navigation. Diverse animals ranging from lobsters to birds are now known to use magnetic positional information for a variety of purposes, including staying on track along migratory pathways, adjusting food intake at appropriate points in a migration, remaining within a suitable oceanic region, and navigating toward specific goals. Recent findings also indicate that sea turtles, salmon, and at least some birds imprint on the magnetic field of their natal area when young and use this information to facilitate return as adults, a process that may underlie long-distance natal homing (a.k.a. natal philopatry) in many species. Despite recent progress, much remains to be learned about the organization of magnetic maps, how they develop, and how animals use them in navigation.

Long-distance transequatorial navigation using sequential measurements of magnetic inclination angle.

Diverse taxa use Earth's magnetic field in combination with other sensory modalities to accomplish navigation tasks ranging from local homing to long-distance migration across continents and ocean basins. Several animals have the ability to use the inclination or tilt of magnetic field lines as a component of a magnetic compass sense that can be used to maintain migratory headings. In addition, a few animals are able to distinguish among different inclination angles and, in effect, exploit inclination as a surrogate for latitude. Little is known, however, about the role that magnetic inclination plays in guiding long-distance migrations. In this paper, we use an agent-based modelling approach to investigate whether an artificial agent can successfully execute a series of transequatorial migrations by using sequential measurements of magnetic inclination. The agent was tested with multiple navigation strategies in both present-day and reversed magnetic fields. The findings (i) demonstrate that sequential inclination measurements can enable migrations between the northern and southern hemispheres, and (ii) demonstrate that an inclination-based strategy can tolerate a reversed magnetic field, which could be useful in the development of autonomous engineered systems that must be robust to magnetic field changes. The findings also appear to be consistent with the results of some animal navigation experiments, although whether any animal exploits a strategy of using sequential measurements of inclination remains unknown.

LACTIC ACIDOSIS INDUCED BY MANUAL RESTRAINT FOR HEALTH EVALUATION AND COMPARISON OF TWO POINT-OF-CARE ANALYZERS IN HEALTHY LOGGERHEAD SEA TURTLES (CARETTA CARETTA).

Sea turtles are often restrained manually for brief periods during veterinary evaluation and care in rescue, rehabilitation, research, and aquarium settings. Blood gas values and lactate are routinely evaluated during triage of sea turtles, and lactate clearance is of prognostic significance in cold-stunned individuals. Although increases in blood lactate have been associated with muscle exertion, experimental forced submergence, trawl and pound net capture, and general anesthesia, changes in blood lactate associated with short periods of manual restraint have not been evaluated. Venous blood gas and lactate values were tested in 16 juvenile loggerhead sea turtles (Caretta caretta) before and after manual restraint for a 15-min routine veterinary examination. The agreement of blood lactate values between two point-of care analyzers (i-STAT and Lactate Plus) was also compared. Blood pH and bicarbonate (HCO3-) decreased significantly (P < 0.001), and partial pressure of carbon dioxide (pCO2) increased significantly (P < 0.0001) after 15 min. Lactate increased significantly between time points for both analyzers (P < 0.0001). Linear regression analysis showed excellent correlation for lactate measurements obtained on both analyzers (r = 0.998). The mean difference in lactate concentrations between the analyzers was statistically significant, indicating that the methods cannot be used interchangeably (P < 0.0001). Deming regression and Bland-Altman plots identified a slight negative proportional bias for lactate measurement by the Lactate Plus compared with the i-STAT. These results suggest that clinicians should evaluate blood gas values and lactate at the beginning of health evaluations and interpret serial lactate values in sea turtles with caution, because even short periods of manual restraint can induce lactic acidosis and considerably influence these values.

Effective mydriasis in juvenile loggerhead turtles (Caretta caretta) following topical administration of rocuronium bromide and 10% phenylephrine.

OBJECTIVE: To determine the combined mydriatic effects of topical rocuronium bromide and phenylephrine in juvenile loggerhead turtles and identify any adverse effects associated with treatment. ANIMALS STUDIED: Eleven juvenile loggerhead turtles (Caretta caretta). PROCEDURES: Four 20 µL drops of rocuronium bromide and four 20 µL drops of 10% phenylephrine were placed into the right eye at 2-minute intervals of 5 turtles, while the same volume of saline was administered to six control turtles. A pupilometer recorded pupil measurements at rest and following a light stimulus at 2, 15, 30, 60, 120, 150, 180, 210, 240, 300, and 360 minutes following delivery of the final drop to the ocular surface. Intraocular pressure (IOP) was also measured at similar time points. RESULTS: The nonilluminated and light-stimulated pupillary diameter of the right eye of treated turtles was significantly greater than baseline starting at 120 and 15 minutes, respectively. Light-stimulated pupillary diameter of treated eyes was greater than that of control eyes from time 15 minutes until the end of the treatment period. No systemic side effects were noted over a 24 hours period following treatment and all turtles showed normal behavior and appetite. No mydriasis was noted in either eye at 24 hours and the anterior segment was normal. CONCLUSIONS: A combination of topical ophthalmic rocuronium bromide and 10% phenylephrine is safe and effective for mydriasis in juvenile loggerhead turtles.

There and back again: natal homing by magnetic navigation in sea turtles and salmon.

Diverse marine animals migrate across vast expanses of seemingly featureless ocean before returning as adults to reproduce in the area where they originated. How animals accomplish such feats of natal homing is an enduring mystery. Growing evidence suggests, however, that sea turtles and salmon imprint on the magnetic field of their home area when young and then use this information to return as adults. Both turtles and salmon have the sensory abilities needed to detect the unique 'magnetic signature' of a coastal area. Analyses have revealed that, for both groups of animals, subtle changes in the geomagnetic field of the home region are correlated with changes in natal homing behavior. In turtles, a relationship between population genetic structure and the magnetic fields that exist at nesting beaches has also been detected, consistent with the hypothesis that turtles recognize their natal areas on the basis of magnetic cues. Salmon likely use a biphasic navigational strategy in which magnetic cues guide fish through the open sea and into the proximity of the home river where chemical cues allow completion of the spawning migration. Similarly, turtles may also exploit local cues to help pinpoint nesting areas once they have arrived in the vicinity. Throughout most of the natal homing migration, however, magnetic navigation appears to be the primary mode of long-distance guidance in both sea turtles and salmon.

Multi-Modal Homing in Sea Turtles: Modeling Dual Use of Geomagnetic and Chemical Cues in Island-Finding.

Sea turtles are capable of navigating across large expanses of ocean to arrive at remote islands for nesting, but how they do so has remained enigmatic. An interesting example involves green turtles (Chelonia mydas) that nest on Ascension Island, a tiny land mass located approximately 2000 km from the turtles' foraging grounds along the coast of Brazil. Sensory cues that turtles are known to detect, and which might hypothetically be used to help locate Ascension Island, include the geomagnetic field, airborne odorants, and waterborne odorants. One possibility is that turtles use magnetic cues to arrive in the vicinity of the island, then use chemical cues to pinpoint its location. As a first step toward investigating this hypothesis, we used oceanic, atmospheric, and geomagnetic models to assess whether magnetic and chemical cues might plausibly be used by turtles to locate Ascension Island. Results suggest that waterborne and airborne odorants alone are insufficient to guide turtles from Brazil to Ascension, but might permit localization of the island once turtles arrive in its vicinity. By contrast, magnetic cues might lead turtles into the vicinity of the island, but would not typically permit its localization because the field shifts gradually over time. Simulations reveal, however, that the sequential use of magnetic and chemical cues can potentially provide a robust navigational strategy for locating Ascension Island. Specifically, one strategy that appears viable is following a magnetic isoline into the vicinity of Ascension Island until an odor plume emanating from the island is encountered, after which turtles might either: (1) initiate a search strategy; or (2) follow the plume to its island source. These findings are consistent with the hypothesis that sea turtles, and perhaps other marine animals, use a multi-modal navigational strategy for locating remote islands.

The magnetic map of hatchling loggerhead sea turtles.

Young loggerhead sea turtles (Caretta caretta) from eastern Florida, U.S.A., undertake a transoceanic migration in which they gradually circle the North Atlantic Ocean before returning to the North American coast. Hatchlings in the open sea are guided at least partly by a 'magnetic map' in which regional magnetic fields function as navigational markers and elicit changes in swimming direction at crucial locations along the migratory route. The magnetic map exists in turtles that have never migrated and thus appears to be inherited. Turtles derive both longitudinal and latitudinal information from the Earth's field, most likely by exploiting unique combinations of field inclination and intensity that occur in different geographic areas. Similar mechanisms may function in the migrations of diverse animals.

Longitude perception and bicoordinate magnetic maps in sea turtles.

Long-distance animal migrants often navigate in ways that imply an awareness of both latitude and longitude. Although several species are known to use magnetic cues as a surrogate for latitude, it is not known how any animal perceives longitude. Magnetic parameters appear to be unpromising as longitudinal markers because they typically vary more in a north-south rather than an east-west direction. Here we report, however, that hatchling loggerhead sea turtles (Caretta caretta) from Florida, USA, when exposed to magnetic fields that exist at two locations with the same latitude but on opposite sides of the Atlantic Ocean, responded by swimming in different directions that would, in each case, help them advance along their circular migratory route. The results demonstrate for the first time that longitude can be encoded into the magnetic positioning system of a migratory animal. Because turtles also assess north-south position magnetically, the findings imply that loggerheads have a navigational system that exploits the Earth's magnetic field as a kind of bicoordinate magnetic map from which both longitudinal and latitudinal information can be extracted.

Geomagnetic imprinting: A unifying hypothesis of long-distance natal homing in salmon and sea turtles.

Several marine animals, including salmon and sea turtles, disperse across vast expanses of ocean before returning as adults to their natal areas to reproduce. How animals accomplish such feats of natal homing has remained an enduring mystery. Salmon are known to use chemical cues to identify their home rivers at the end of spawning migrations. Such cues, however, do not extend far enough into the ocean to guide migratory movements that begin in open-sea locations hundreds or thousands of kilometers away. Similarly, how sea turtles reach their nesting areas from distant sites is unknown. However, both salmon and sea turtles detect the magnetic field of the Earth and use it as a directional cue. In addition, sea turtles derive positional information from two magnetic elements (inclination angle and intensity) that vary predictably across the globe and endow different geographic areas with unique magnetic signatures. Here we propose that salmon and sea turtles imprint on the magnetic field of their natal areas and later use this information to direct natal homing. This novel hypothesis provides the first plausible explanation for how marine animals can navigate to natal areas from distant oceanic locations. The hypothesis appears to be compatible with present and recent rates of field change (secular variation); one implication, however, is that unusually rapid changes in the Earth's field, as occasionally occur during geomagnetic polarity reversals, may affect ecological processes by disrupting natal homing, resulting in widespread colonization events and changes in population structure.

The sensory ecology of ocean navigation.

How animals guide themselves across vast expanses of open ocean, sometimes to specific geographic areas, has remained an enduring mystery of behavioral biology. In this review we briefly contrast underwater oceanic navigation with terrestrial navigation and summarize the advantages and constraints of different approaches used to analyze animal navigation in the sea. In addition, we highlight studies and techniques that have begun to unravel the sensory cues that underlie navigation in sea turtles, salmon and other ocean migrants. Environmental signals of importance include geomagnetic, chemical and hydrodynamic cues, perhaps supplemented in some cases by celestial cues or other sources of information that remain to be discovered. An interesting similarity between sea turtles and salmon is that both have been hypothesized to complete long-distance reproductive migrations using navigational systems composed of two different suites of mechanisms that function sequentially over different spatial scales. The basic organization of navigation in these two groups of animals may be functionally similar, and perhaps also representative of other long-distance ocean navigators.

Magnetic maps in animals: nature's GPS.

Diverse animals detect the Earth's magnetic field and use it as a cue in orientation and navigation. Most research on magnetoreception has focused on the directional or ;compass' information that can be extracted from the Earth's field. Because the field varies predictably across the surface of the globe, however, it also provides a potential source of positional or 'map' information, which some animals use to steer themselves along migratory pathways or to navigate toward specific target areas. The use of magnetic positional information has been demonstrated in several diverse animals including sea turtles, spiny lobsters, newts and birds, suggesting that such systems are phylogenetically widespread and can function over a wide range of spatial scales. These ;magnetic maps' have not yet been fully characterized. They may be organized in several fundamentally different ways, some of which bear little resemblance to human maps, and they may also be used in conjunction with unconventional navigational strategies.

Animal behaviour: geomagnetic map used in sea-turtle navigation.

Migratory animals capable of navigating to a specific destination, and of compensating for an artificial displacement into unfamiliar territory, are thought to have a compass for maintaining their direction of travel and a map sense that enables them to know their location relative to their destination. Compasses are based on environmental cues such as the stars, the Sun, skylight polarization and magnetism, but little is known about the sensory mechanism responsible for the map sense. Here we show that the green sea-turtle (Chelonia mydas) has a map that is at least partly based on geomagnetic cues.