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Light-Induced Changes in the Prey Consumption and Behavior of Two Juvenile Planktivorous Fish

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Light-Induced Changes in the Prey Consumption and Behavior of Two Juvenile Planktivorous Fish 1 MARINE ECOLOGY PROGRESS SERIES Vol. 181: 41-51,1999 Published May 18 1 Mar Ecol Prog Ser Light-induced changes in the prey consumption and behavior of two juvenile planktivorous fish Clifford H. Ryer*, Bori L. Olla Fisheries Behavioral Ecology Group, Alaska Fisheries Science Center, National Marine Fisheries Service, NOAA. Hatfield Marine Science Center, Newport. Oregon 97365. USA ABSTRACT: Walleye pollock and sablefish, as O+ yr juveniles, are pelagic particulate feeding plankti- vores. We conducted a series of laboratory experiments to determine how illumination influences prey consumption in these species, utilizing live Artemia sp. as prey. Both juvenile walleye pollock and sablefish were characterized by a sigmoidal relationship between the log of illumination and the num- ber of prey consumed, with greater prey consumption at higher illuminations. The threshold illumina- tion below which fish were no longer able to visually forage was approximately 5 X 10-' pE S-' m-* for walleye pollock and 5 X IO-~pE S-' me2for sablefish, indicating that walleye pollock are better adapted for visual feedng at depth or at night than are sablefish. This is consistent with what is known about their vertical distributions at this life stage; walleye pollock make daily vertical migrations which keep them at lower illuminations than sablefish, which remain at or near the water surface throughout the die1 cycle. Although feeding more effectively in the light, both species were capable of detecting and capturing prey in darkness. KEY WORDS: Visual threshold . Nonvisual feeding behav~or. Illumination . Nocturnal INTRODUCTION may vary dramatically on an hourly, daily, monthly and seasonal basis. Although some fishes avoid bright light Many planktivorous fish are visual foragers, highly (e.g.Olla & Davis 1990, Sogard & Olla 1993), the abil- dependent upon light to efficiently detect and con- ity to visually detect prey is probably more often lim- sume their zooplankton prey. Therefore, understand- ited by conditions of low illumination. As a result, a ing the role of light in determining where and when number of studies have examined the illumination these fish feed is a necessary underpinning to model- thresholds for feeding in various fishes, with the goal of ing their energetics, growth and survival. The quantity predicting the conditions under which they will be and quality of light available in aquatic environments able to forage (Blaxter 1965, Hunter 1968, Bagarinao & is determined, first of all, by astronon~icaland meteo- Hunter 1983, Dabrowski & Jewson 1984, Bergman rological conditions. At the water surface, illumination 1988, Connaughton et al. 1994, Huse 1994, McMahon may span 9 orders of magnitude, from full daylight & Holanov 1995). For example, Hunter (1968) found with clear skies to moonless nights under dark storm that jack mackerel Trachurus symmetricus feed effec- cloud conditions (US.Department of the Navy 1952). tively on Artemia sp. down to 6 X 10-5 ft-L (approxi- Secondly, light will be further reduced by unfavorable mately 1 X 10-= pE S-' m-'), suggesting that they may sea surface conditions, turbidity and many other water be capable of feeding in surface waters at night when quality variables which may vary with depth and affect there is moonlight. spectral attenuation (McFarland 1986). Thus, the opti- The light requirements for feeding by visual plankti- cal environment in which these fish live and forage vores may also provide insights into other aspects of fish ecology. For example, larval and juvenile fish often undergo daily vertical migrations, with the most com- 0 lnter-Research 1999 Resale of full article not permitted 4 2 Mar Ecol Prog Ser 181: 41-51, 1999 mon pattern involving upward movement to surface MATERIALS AND METHODS waters at or near dusk, followed by a return to the depths at dawn (Neilson & Perry 1990). Whatever the Fish collection and maintenance. During June 1996, causes and proximal mechanism involved in vertical juvenile walleye pollock were collected at night from migration, it unquestionably influences the visual Puget Sound, Port Townsend, WA, USA, by dipnetting environment in which fish must forage, and, con- them as they aggregated around a light. Juvenile versely, it is probable that visual capabilities may par- sablefish Anaplopoma fimbria were captured during tially dictate the range of vertical movements under- April 1996 using a neuston net at night, approximately taken. 32 km off Newport, OR, USA. At the time of collection, In this study, we examine the role of light in the for- juveniles ranged from 20 to 50 mm TL (total length). aging of 2 pelagic planktivorous fishes: juvenile wall- Fish were transported back to our laboratory in New- eye pollock Theragra chalcogramma and juvenile port and maintained in 3000 1 circular tanks provided sablefish Anaplopoma fimbria. Both species are inhab- with a continuous flow of seawater: salinity range 28 to itants of the coastal waters of the north Pacific and 33%o,temperature range 9 to 13°C. Both species were form the basis of important commercial fisheries. As fed pelletized food daily, and, while the exact ration juveniles, walleye pollock often migrate vertically, was not measured, it was sufficient to promote growth. ascending at dusk and descending at dawn (Bailey Fish were held for 2 to 4 wk prior to experimentation. 1989). It is unknown whether this migration is a Although they received no live food during this period, response to predation pressure, feeding opportunities, preliminary experiments revealed that both walleye energetic considerations, or a combination of these fac- pollock and sablefish would readily attack live prey if tors, but it clearly narrows the range of light intensities given the opportunity. experienced on a daily basis, compared, that is, to a Infrared observation tanks. Experiments were con- fish that exclusively occupies surface waters. Juvenile ducted using 4 identical glass tanks (50 X 50 X 50 cm, walleye pollock have relatively large eyes and have W X L X H) supported by a plexiglass platform and sur- generally been assumed to be visual foragers (Olla & rounded by a light-proof blind in a dark room. A single Davis 1990).Juvenile walleye pollock decrease activity inflow pipe, which extended vertically to within 1 cm at night (Sogard & Olla 1996, Ryer & Olla 1998) a.nd of the tank bottom, supplied seawater to each tank, cease schooling (Brodeur & Wilson 1996, Ryer & Olla while 2 drain holes allowed for outflow. For each tank, 1998). These observations suggest that the walleye a feeding tube passed through a sleeve in the blind, pollock is a visually oriented diurnal species, which allowing the tube to be lowered to the water's surface has been reinforced by behavioral studies demonstrat- to introduce prey and then withdrawn with minimal ing that they visually monitor foraging conspecifics for disturbance. Each tank was illuminated from below cues that may reveal the location of prey patches with a 60 W LED infrared (IR) illuminator. The wave- (Baird et al. 1991, Ryer & Olla 1995, 1997). This does length of emitted IR light peaked at 880 nm, with no not preclude, however, their being capable of feeding emissions below 760 nm. Fish are insensitive to light in under conditions of low light by using sensory modali- this range (Douglas & Hawryshyn 1990) and IR illumi- ties other than vision (e.g.Merati & Brodeur 1996). nation has been utilized by other researchers to exam- Juvenile sablefish also school, but are more active Ine fish behavior under low visible light conditions than walleye pollock (Ryer & Olla 1997).Like the wall- (John 1964, Pitcher & Turner 1986, Higgs & Fuiman eye pollock, they also feed upon zooplankton, but 1996). A sheet of light diffusing material, commonly available data suggest that they do not make extensive utilized in overhead fluorescent fixtures, was posi- daily vertical migrations, remaining, instead, near the tioned between the tanks and the plexiglass platform, surface both day and night (Shenker & Olla 1986, Sog- and, when ~lluminatedfrom below, produced a diffuse ard & Olla 1998). In addition, these data indicate that bright background against which fish could be viewed juvenile sablefish are diurnal feeders, suggesting that as silhouettes from above. A video camera, with spec- they may not be capable of detecting prey at low illu- tral sensitivity well into the IR range, was positioned minations (Sogard unpubl.). above each tank and cabled to a video recorder and We exposed both species to surrogate planktonic monitor in another room. Four green LED illumi.nators prey, live Artemia sp., under varying illumination, were positioned above each tank to provide visible determining the number of prey consumed and ob- illumination. These illuminators emitted within a nar- serving foraging behavior. On the basis of what was row spectral range of approximately 10 nm, with a known about their vertical distributions and feeding peak at 555 nm, and were digitally controlled using a behavior, we hypothesized that juvenile walleye pol- micro-computer, allowing incremental manipulation of lock would have a lower light threshold than juvenile illumination from 1.90 X 10-' to 1.55 X IO-~pE S-' m-', sablefish for visually mediated foraging. measured at the water surface. Higher light levels Ryer & OUa: Influence of light on planktivorous fish 43 (1.50 X and 1.50 X 10-' pE S-' m-') were achieved which they were videotaped during a 30 rnin pre-feed- by augmenting the green LED output with the addi- ing period, followed by prey introduction and video- tional light emitted by a rheostat controlled 15 W in- taping of feeding behavior. Live adult Arternia sp., candescent bulb positioned above each tank. obtained weekly from a pet store, were used as prey. Light measurements. Measurements in each repli- Prey were not fed during the study. Two hundred prey cate tank were made at the beginning of the study, but were counted and placed into a 100 m1 beaker of sea- occasional measurements at later dates confirmed that water 30 rnin prior to being introduced to the IR tank.
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