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JOBLING ET AL. REVIEWS IN FISHERIES SCIENCE <br />V. D/EL FEEDING PATTERNS <br />In addition to having application in the estimation of daily feed intake, the <br />X-radiographic method allows investigation of diel changes in feeding behavior. Diel <br />feeding patterns of Arctic charr have been examined at different seasons of the year <br />for fish taken from the wild (J~rgensen and Jobling, 1989), and for those reared in <br />a hatchery environment throughout their lives. Marked seasonal variations were <br />recorded in the times of day during which fish fed most actively (Figure 2), but <br />patterns of feed intake were similar for the wild-caught and hatchery-reared charr. <br />At certain times of the year, the charr consumed a large proportion of their daily feed <br />intake at night, and there were usually peaks of feeding activity around dusk and <br />dawn. The overall diel and seasonal changes in feeding patterns of Arctic charr <br />appear to be similar to those of rainbow trout (Oncorhynchus mykiss) (Landless, <br />1976), but differ markedly from those of Atlantic salmon, which appear to feed <br />predominantly during daylight hours, irrespective of season (Higgins and Talbot, <br />1985; Jorgensen and Jobling, 1992). <br />Although the X-radiographic technique can be used to study diel changes in <br />patterns of feed intake (Higgins and Talbot, 1985; Jorgensen and Jobling, 1989, <br />1992), information about feeding activity is collected more easily and cheaply using <br />on-demand feeding systems (Landless, 1976; Alanara, 1992a,b; Boujard et ai., 1992). <br />On-demand feeding systems may be constructed to possibly allow monitoring each <br />release of feed as a discrete event under conditions where feed is continuously <br />available to the fish. Thus, these systems enable long-term automatic recording of <br />data without disturbance to the fish. <br />Vl. MODES OF FEEDING BEHAVIOR <br />The fact that Arctic charr often consumed much of their daily feed intake at night <br />Qorgensen and Jobling, 1989) led to the hypothesis that foraging on the bottom may <br />be an important feeding mode for the species. The hypothesis was tested by <br />comparing the daily feed intakes of groups of fish prevented from reaching feed <br />lying on the tank bottom with intakes of fish allowed free access to the bottom. <br />Irrespective of the lighting conditions used in the tests, the feed intake of charr <br />allowed access to feed lying on the tank bottom was significantly greater than intake <br />of charr forced to forage in the water column (Jorgensen and Jobling, 1990). When <br />a similar study was carried out on Atlantic salmon (Jorgensen and Jobling, 1992), the <br />results were different from those obtained in the charr experiments. Atlantic salmon <br />appear to be far less dependent on bottom foraging in order to obtain adequate <br />amounts of feed (Figure 3). <br />The results presented above refer to short-term experiments, and further con- <br />firmation of the importance of bottom feeding for charr was obtained in longer-term <br />growth experiments conducted over a period of 17 weeks. On each of the four <br />occasions on which daily feed intake was estimated, fish allowed access to feed lying <br />on the bottom had consumed significantly more than the fish forced to forage in the <br />water column. These differences in feed intake affected growth. The Arctic charr <br />given free access to the feed lying on the tank bottom almost doubled their body <br />230 <br />