7

A History of Telemetry in Fishery Research

Eric E. HockErsmitH and JoHn W. BEEman

Section 2

Biotelemetry has been defined as “the instrumental technique for gaining and transmitting information from a living organism and its environment to a remote observer” (Slater 1965). Biotelemetry typically utilizes wireless transmission of either an audible signal or electronic data to determine location of a tagged animal. Fisheries researchers use location information to gain a variety of insights into migration, habitat use, behavior, productivity, or survival of fish. Biotelemetry can be divided into two basic categories, acoustic or radio, based on mode of transmission, mechanical or electromagnetic energy, and operating frequency. Most acous-tic systems in use today transmit at low frequency, between 30 and 300 kHz, while most radio systems transmit at very high frequency, between 30 and 300 MHz (Sisak and Lotimer 1998).

Acoustic telemetry is based on the principals of sonar (sound navigation and ranging), which was developed to detect submarines during World War I. The properties of acoustic systems favor their use in deep waters with high conductivity and low turbulence (Winter 1996). Radio telemetry is based on the principals of wireless radio communication, which were first demonstrated by Nikola Tesla in 1893. Radio systems are best suited in shallow wa-ters with relatively low conductivity but have the added benefit of improved signal detection in turbulent conditions and with aerial antennas. Advances in both technologies have resulted in highly efficient transmitter and receiving systems.

Advancements in products used for animal telemetry over the past 50 years have gener-ally followed those in the electronics field (Figure 1). Bell Laboratories1 ushered in the age of digital electronics with the invention of the transistor in 1947 (Mann 2000). Today transis-tors are common in everyday items such as radios, televisions, hearing aids, computers, cell phones and even MP3 players. Consumer demand for inexpensive small electronic devices with increased functionality has continually driven advancements in the field of electronics. These advancements have subsequently led to improvements in biotelemetry transmitters and receivers such as miniaturization of components, increased battery performance, and more powerful micro-processing.

The earliest use of biotelemetry to study fish and wildlife occurred in the mid- to late 1950s. Research covered a broad field, including examination of salmon migrations (Tre-fethen 1956), measurement of incubation temperatures for penguin eggs (Ecklund and Charl-tin 1959), and investigation of woodchuck movements (LeMunyan et al. 1959). Telemetry brought two new advantages to fish and wildlife research: the ability to identify individual 1Reference to trade names does not imply endorsement by the U.S. Government.

8 Section 2

animals, and the ability to locate individual animals without having to observe or recapture them, greatly enhancing behavior and migration studies. Further, the quantity and detail of data generated from studies using biotelemetry has often been greater than that generated from mark–recapture or direct observation methods (Adams and Davis 1967).

The first acoustic telemetry equipment for studying fish was developed in 1956 by the U.S. Bureau of Commercial Fisheries (BCF) and the Minneapolis-Honeywell Regulator Cor-poration (Figure 2; Trefethen 1956; Trefethen et al. 1957). Fish were externally tagged with a transmitter attached to the dorsal area of the fish (Figure 3). These studies examined adult salmon passage behavior at Bonneville Dam on the Columbia River (Trefethen 1956; John-son 1960). Fish locations and movements were determined by mobile tracking from a boat (Figure 4) and using fixed-site acoustic receivers that recorded the data on paper and magnetic tape (Figure 5). In considering the advances in knowledge of fish movement and migration, Arnold (2000) concluded that the insight gained during the 30 years following application of biotelemetry for fisheries research were greater than those during the preceding 75 years.

Acoustic telemetry use in fisheries research continued to develop during the 1960s, fol-lowing invention of the integrated circuit by Texas Instruments in 1958. Because of their size, early acoustic transmitters could be used only with large adult fish. However, within a few years of initial development, an acoustic tag was produced that was approximately 25% smaller and had twice the detection range and three times the tag life of earlier designs (Novotny and Esterberg 1962). At approximately the same time, University of Wisconsin researchers developed a much smaller acoustic transmitter for use with white bass. These fish had an average length of only 300 mm, a size considerably less than adult Chinook salmon Oncorhynchus tshawytscha, but due to battery limitations, the tag had a life of less than 24 h (Henderson et al. 1966).

In the early 1960s, the BCF used acoustic-tagged fish to determine whether the newly formed Brownlee Reservoir on the Snake River impacted the ability of adult Chinook salmon to migrate to traditional spawning grounds (Trefethen 1968). They used both mobile tracking and fixed-site monitoring to locate tagged fish. Similar equipment was used to study Chinook

Figure 1. Time line of major developments in electronics that contributed to advances in fisheries telemetry over the past 200 years. Telemetry specific developments are highlighted.

9A History of Telemetry in Fishery Research

Figure 2. The first acoustic receiver and transmitter developed to study migration and behavior of adult salmon in the Columbia River (Trefethen 1956; Trefethen et al. 1957).

10 Section 2

Figure 3. Early acoustic tag which used a hog ring clamped into the dorsal muscle of the fish for attach-ment.

Figure 4. Acoustic tracking system used for mobile tracking adult salmon from a boat during the late 1950s near Bonneville Dam on the Columbia River.

11A History of Telemetry in Fishery Research

salmon movements through the San Joaquin River delta in California in 1964 (Hallock et al. 1970). In the USSR, acoustic transmitters developed in 1964 were used to study sturgeon mi-grations in 1965 (Malinin and Svirskii 1973). By 1967, acoustic telemetry was used routinely, and applications were expanded to investigation of more fine-scale fish movement, such as a study of net avoidance by American shad in the Connecticut River (Leggett and Jones 1971).

During the late 1960s and early 1970s, research programs such as those at the Marine Biotelemetry Laboratory at the University of New Brunswick led to improvements in acous-tic signal transmission efficiency in water from about 1–80%. The ability to automatically receive and decode transmitted information was also developed around this time (D. Pincock, Electrical Engineering Department, Technical University of Nova Scotia, personal communi-cation), along with the ability to transmit pressure and temperature information from acoustic tags (Luke et al. 1973; Rochelle and Coutant 1973). Many transmitters used in these programs were still hand-built from analog parts soldered together under stereoscopes. The increasing availability of digital rather than analog equipment during the 1970s enabled more efficient construction techniques and products that were more reliable.

Radio telemetry was first used by wildlife biologists to study movement of terrestrial animals in 1956 (LeMunyan et al. 1959), and its use grew rapidly during the 1960s (Cochran and Lord 1963). However, this technology was not used to study fish until 1968 (Lonsdale and Baxter 1968). During the 1970s, a variety of researchers began modifying radio telemetry systems used in terrestrial habitats for use in freshwater.

As the use of acoustic telemetry increased, fisheries researchers found that it performed poorly in some environments, such as areas of high turbulence (Trefethen 1956; Johnson 1960) or large areas with limited access for monitoring. In 1971, National Marine Fisheries Service (formerly BCF) researchers switched from acoustic to radio tags to evaluate the upstream migration behavior of Chinook salmon. Radio tags were chosen because they performed well

Figure 5. Fixed-site acoustic receiver system used during the late 1950s near Bonneville Dam on the Co-lumbia River. Data was recorded on both paper and magnetic tape.

12 Section 2

in the turbulent conditions below dams and had higher detection rates than acoustic transmit-ters (Monan et al. 1975). The ability to detect radio transmitters in air using highly portable aerial antennas also provided a major advantage for migration studies in large or inaccessible river systems (Koehn 1999). Furthermore, fisheries researchers benefitted significantly by using the methods, equipment, and analytical techniques developed from wildlife telemetry. By the mid-1970s, radio telemetry and its newly discovered advantages had eclipsed acoustic telemetry to the point where use of the latter was limited to systems with high conductivity where radio telemetry worked poorly (Stasko and Pincock 1977).

Prior to about 1970, most researchers using acoustic and radio telemetry made at least some of their own equipment (Lonsdale and Baxter 1968; Standora et al. 1972), or worked with private sector electronic engineers, as in the relationship between the BCF and Honey-well (Trefethen et al. 1957). It was common for fisheries researchers to describe circuit dia-grams of telemetry transmitters when publishing their biological results (Cochran and Lord 1963; Luke et al. 1973). Radio telemetry transmitters and receivers were first made com-mercially available in 1967 by AVM Instrument Company (Figure 6). For acoustic telemetry, commercial equipment was first produced by Sonotronics, founded in 1971. During the early 1970s acoustic receivers were also developed by the Marine Biotelemetry Laboratory at the University of New Brunswick and Smith Root (Figure 7).

Both technological advances and biotelemetry companies have proliferated since the 1970s. Today more than 30 companies produce equipment for biotelemetry applications worldwide, with the majority located in North America. Roughly twice as many manufactur-ers produce equipment for radio telemetry compared to acoustic telemetry, and only a few produce both. The higher prevalence of radio telemetry equipment manufacturers is likely due to greater demand, resulting from the ability of radio telemetry to be used in both aquatic and

Figure 6. Early versions of commercially produced radio telemetry receivers. The receiver on the left was produced by AVM in the late 1960s and the receiver on the right was produced by Smith-Root in the early 1970s.

13A History of Telemetry in Fishery Research

terrestrial settings. About one-third of all biotelemetry startup companies formed during the past 50 years began during the 1980s following the rapid advances in electronic components during this period (Arnold and Dewar 2001).

Commercialization of telemetry equipment allowed the technology to be used by re-searchers who did not have the capability of manufacturing their own equipment. Between 1970 and 1980, the same advances in electronics that led to the first digital cell phones, lithium batteries, and personal computers were used by biotelemetry firms to increase the ca-pability of commercial transmitters and receivers. Data-logging radio and acoustic receivers became more refined and more available during this period, and transmitters continued to be-come smaller and more capable. Reduction in tag size, while either maintaining or improving detection range and tag life, continues to be a primary objective of innovation in transmitter development (Figure 8). Today very small acoustic and radio transmitters are readily avail-able, with some as light as 0.2 g in air (Figure 9; Naef-Daenzer et al. 2005) and have provided opportunities to study species that previously could not be tagged due to their small size, as well as the juvenile life stages of many species.

There are two types of transmitters—noncoded (typically referred to as pinger tags) or coded. Further description and an explanation of transmitters and coding systems is provided in Section 5. The earliest studies used noncoded transmitters because existing receivers were simple in design and did not contain the circuitry to process the signals from coded transmit-ters. In 1982, the National Marine Fisheries Service developed the first pulse code modulation radio transmitter, with a code set that provided the ability to study up to 600 individuals at a time (Stuehrenberg et al. 1990). They used the new transmitters in the Columbia River Basin to study salmonid migration and dam passage, first of adults during 1982, and subsequently of juveniles during 1985. The coding of these transmitters provided the ability to greatly in-crease the number of animals tracked, as well as to decrease the scan time of receivers. By the early 2000s there were several commercial coding schemes available, with most transmitter manufacturers using their own.

Wide-band receivers that could simultaneously scan multiple radio frequencies were de-veloped shortly after the advent of coded transmitters. One of the first wide-band receivers could decode and record up to 144 unique transmitters that simultaneously occupied a single detection field (Stuehrenberg et al. 1990). Prior to the 1990s, most multi-frequency receivers

Figure 7. Acoustic telemetry receivers from the 1970s. The receiver on the left was developed by The Uni-versity of New Brunswick Marine Biotelemetry Lab (photo courtesy of Doug Pincock) and the receiver on the right was produced by Smith-Root.

14 Section 2

Figure 8. Relative size and weight in air of radio transmitters used by National Marine Fisheries Service to study juvenile salmon migration in the Columbia River over a 27-year period from 1980 through 2007.

Figure 9. Circuit schematic (A) and photograph (B) of a 0.2 g radio transmitter (Naef-Daenzer, et al. 2005).

15A History of Telemetry in Fishery Research

either mechanically or automatically stepped sequentially through each frequency or antenna, one at a time. Most wide-band receivers today are based on digital spectrum processors, which were first developed in the late 1970s. They virtually eliminate the scan time required by narrow-band receivers, but are less sensitive to weak transmissions. Wide-band receivers are available from several manufacturers today and are particularly advantageous for detect-ing tags in applications where fish could move through a detection field rapidly.

Although the first artificial satellite was launched in 1957, satellites were not used in conjunction with fish and wildlife telemetry until the early 1970s. The first integration of satellite technology and telemetry was used in a study tracking radio-tagged bears (Buech-ner et al. 1971). In 1978, a worldwide tracking and environmental monitoring system was started through a collaborative relationship between the French Space Agency, U.S. National Aeronautics and Space Administration, and the U.S. National Oceanic and Atmospheric Ad-ministration. This partnership led to the Advanced Research and Global Observation Satellite (ARGOS) system that used polar-orbiting satellites and up to 50 ground stations to collect data from more than 20,000 ocean platforms. ARGOS was originally developed for meteo-rological and oceanographic applications, but in the 1980s, transmitter miniaturization com-bined with an effective data logger, led to applications for tracking fish. Frequencies used to track fish are low compared to those used for satellite communications. The ground stations and floating platforms are used to collect data which is then converted to the frequency used by the satellites for data transmission.

The Global Positioning System (GPS) was developed in the 1970s to provide three-di-mensional positioning for military applications using satellites (Musser 1992). Applications of GPS have been used with aquatic telemetry to describe locations of radio-tagged fish mi-grating through large river systems (Hockersmith and Peterson 1997) and for tasks as simple as synchronizing clocks in a series of telemetry receivers. Satellite systems have also been used for retrieval of data from receivers (Eiler 1995), or the tags themselves (Block et al. 1998), thus expanding the possible applications in remote areas.

In addition to the traditional methods, several telemetry applications using specialized tags have been developed for fisheries research. Some of these include data archival and geo-locating archival transmitters; the CART, or combined acoustic and radio transmitter (Solomon and Potter 1988); the PSATS, or pop-up satellite archival tag (Block et al. 1998); the SPOT, or smart position transmitting tag; and the CHAT, or communicating histogram acoustic transponder tag (Voegeli et al. 2001).

Researchers have either combined sensors with telemetry or used telemetry to relay data from sensors since 1957 (Eklund and Charlton 1959). The first description of sensor use and telemetry in the aquatic environment was the measurement of body temperature for an un-tethered dolphin (Mackay 1964). Sensor technologies have been combined with both acoustic and radio transmitters to measure either the environment occupied by or the physiology of the tagged animal, along with its location.

In fish telemetry studies that measure environmental conditions, typical sensors used in-clude depth (Beeman et al. 1998), temperature (Haynes et al. 1986), and ambient light (Stan-dora et al. 1972). Tilt sensors have been used to indicate activity and mortality in telemetry studies (Eiler 1990). Early versions of these used a mercury switch; however these have been replaced by switches that incorporate a metal rolling ball. Physiological studies of tagged fish have primarily used electromyogram (EMG) transmitters, which measure the electrical activ-ity of muscles from electrodes inserted into the musculature of the fish (Kaseloo et al. 1991).

16 Section 2

EMG has been used to measure swimming speed and to estimate energetic costs associated with physical activity such as movement through a fishway at a dam (Geist et al. 2000). EMG transmitters can also be used to measure mortality, since muscle activity ceases at death.

The field of specialized transmitters and their application continues to expand rapidly with growth in the electronics industry. This expansion is reflected in the prevalence of telem-etry publications, which has increased greatly over the past 50 years. For example, in 1972, a review of underwater biotelemetry examined 52 papers (Malinin and Svirskii 1973). Five years later, a similar review was comprised of 147 publications (Stasko and Pincock 1977), and fourteen years later, a review by Baras (1991) included over 1,100 references. Other biotelemetry reviews include those completed by Ireland and Kanwisher (1978) and Mitson (1978).

Expansion of biotelemetry applications was also reflected in the appearance of several conference series specific to the field of biotelemetry. The longest running of these was or-ganized by the International Society of Biotelemetry (ISOB; www.biotelmetery.org), which held 17 conferences between 1974 and 2003. The ISOB was founded in 1973 to encourage research in biotelemetry and related fields, and to promote its application in medicine and life sciences. More recently, ISOB requests for presenters have seen a poor response, and no conferences were held since 2003 (J. Eiler, National Marine Fisheries Service, personal communication). Another long-term series was the International Conference on Wildlife Bio-telemetry, first held in Laramie, WY, in 1977. This conference was held every two years until the last meeting in 1985. The Conference on Fish Telemetry in Europe began meeting every other year in 1995 and is still well attended today.

These specialized meetings provided a vital mechanism for the development of technical aspects and applications in an emerging field prior to the startup of the commercial sector. They provided a venue for researchers to demonstrate advances in technology and commer-cial vendors to promote new products. When wildlife and fish telemetry was in its infancy, getting a diverse group of researchers together furthered the development of technical as-pects and applications, particularly since those using the technique were relatively few. As the technology developed, it has become widely accepted by the main stream disciplines (e.g., American Fisheries Society, Wildlife Society, etc.). Today biotelemetry equipment is widely available and its applications are common, with frequent presentations of telemetry studies at fisheries conferences, and conferences in many other fields. The transition from an emerging tool that is continually improving and evolving to one that is commonly used, combined with an increasing shift towards holistic ecosystem studies, has likely shifted interest to specific fields rather than general methodology.

Advances in animal telemetry have opened the door for collaboration among researchers across the globe, and several ambitious projects were underway as of this writing. Below we enumerate some of these applications to show the level of creativity and diversity within the field. However, due to the broad scope and speed of expansion in this field, we do not attempt to provide a comprehensive list or to cover these topics in detail.

One large collaborative project is the Ocean Tracking Network, a multinational effort to understand marine life, ocean conditions, and how the changing earth affects them. It is based on acoustic tracking equipment installed in 14 ocean regions of all seven continents. Transmitters in marine animals transmit data to underwater hydrophones, to one another, and eventually to land-based system via satellite or cables (see OceanTrackingNetwork.org).

17A History of Telemetry in Fishery Research

A second large marine system collaboration using telemetry is the Census of Marine Life. Telemetry projects associated with the census include the Pacific Ocean Shelf Tracking (www.postcomol.org) and Tagging of Pelagic Predators (www.topp.org) projects. The POST project uses a series of listening lines placed across the continental shelf of North America to monitor passing marine animals tagged with compatible technology. The TOPP project uses a variety of animal-locating tags, such as PSAT and SPOT, to study movements of large pelagic animals in the North Pacific Ocean.

Large studies examining extensive migration are also being conducted in freshwater. For example, international studies between the U.S. and Canada used radio telemetry to examine adult salmon migrations over thousands of miles of the Yukon River and provide information on basin-wide stock composition and timing (Eiler et al. 2004). Extensive systems of acoustic receivers have been deployed in freshwater environments such as the Sacramento and Co-lumbia Rivers in the United States. These systems were installed to study factors affecting the migration and survival of juvenile salmonids during their seaward migration, such as passage of hydropower dams or water withdrawal projects.

With continued technological innovation, future applications of telemetry will likely pro-vide insights that are currently unavailable. Improvements in battery technology, combined with continued miniaturization of transmitter components, will likely reduce transmitter size further, while increasing efficiency and extending either detection range or tag life. As trans-mitter circuitry becomes more efficient, surplus battery power can be used to power sensors, so that a common practice of future studies will be to study not only the movement and be-havior of tagged fish but also to simultaneously collect information on the environment fish use. This will lead to more comprehensive multidiscipline studies addressing behavior, biol-ogy, and ecology. In the future, technological advances may eventually lead to a transmitter capable of following the movements and behavior of individuals throughout their life cycle.

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