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Jamming avoidance response

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278:. Ampullary receptors respond to low-frequency stimulation less than 40 Hz and their role in the JAR is currently unknown. Tuberous receptors respond to higher frequencies, firing best near the fish's normal EOD frequency. Tuberous receptors themselves have two types, the T-unit and P-unit. The T-unit (T standing for time, meaning phase in the cycle) fires synchronously with the signal frequency by firing a spike on every cycle of the waveform. P-units (P standing for probability) tend to fire when the amplitude increases and fire less when it decreases. Under conditions of jamming, the P-unit fires on the amplitude peaks of the beat cycle where the two waves constructively interfere. So, a combined stimulus-EOD signal causes T-units to fire at the intermediate frequency, and causes P-unit firing to increase and decrease periodically with the beat. 241: 685: 22: 625: 557: 552: 131: 512: 680: 620: 1774: 3112: 2766: 2748: 517: 448: 453: 2778: 3124: 314:, which then projects onto two different pathways. Neurons selective for a positive difference (stimulus greater than EOD) stimulate the prepacemaker nucleus, while neurons selective for a negative difference (stimulus less than EOD) inhibit the sublemniscal prepacemaker nucleus. Both prepacemaker nuclei send projections to the pacemaker nucleus, which ultimately controls the frequency of the EOD. 302:. Phase and amplitude information are integrated here to determine whether the stimulus frequency is greater or less than the EOD frequency. Sign-selective neurons in the deeper layers of the torus semicircularis are selective to whether the frequency difference is positive or negative; any given sign-selective cell fires in one case but not in the other. 202:
To determine how close the stimulus frequency is to the discharge frequency, the fish compares the two frequencies using its electroreceptive organs, rather than comparing the discharge frequency to an internal pacemaker; in other words, the JAR relies only on sensory information. This was determined
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called spherical cells in the electrosensory lateral line lobe. By combining information from multiple T-units, the spherical cell is even more precise in its time coding. Amplitude-coding P-units converge onto pyramidal cells, also in the electrosensory lateral line lobe. Two types of pyramidal
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typically are within the electric field range of three to five other fish of the same species at any time. If many fish are located near each other, it is beneficial for each fish to distinguish between their own signal and those of others; this can be done by increasing the frequency difference
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near the fish, if the stimulus frequency is within 5 Hz of the fish's electric organ discharge (EOD) frequency, the fish alters its EOD frequency to increase the difference between its own frequency and the stimulus frequency. Stimuli above the fish's EOD frequency push the EOD frequency
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most likely arose independently in the two lineages. Weakly electric fish are mostly pulse-dischargers, which do not perform the JAR, while some are wave-dischargers. Wave-discharge evolved in two taxa: the superfamily Apteronotoidea (order
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to create electric fields, and they detect small distortions of these fields using special electroreceptive organs in the skin. All fish with the JAR are wave-discharging fish that emit steady quasi-sinusoidal discharges. For the genus
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If a neighboring sinusoidal electric field is discharging close to the fish's EOD frequency, it causes interference which results in sensory confusion in the fish and sufficient jamming to prevent it from electrolocating effectively.
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downwards, while frequencies below that of the fish push the EOD frequency upwards, with a maximum change of about ±6.5 Hz. This behavior was given the name "jamming avoidance response" several years later in 1972, in a paper by
207:, and then stimulating the fish with two external frequencies. The JAR, measured from the electromotor neurons in the spinal cord, depended only on the frequencies of the external stimuli, and not on the frequency of the pacemaker. 247:
sensory coding for jamming avoidance response. The fish avoids interference with its electrolocation signal by changing its frequency when a jamming signal is detected. This is mediated by a T-unit
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cells exist: excitatory E-units, which fire more when stimulated by P-units, and inhibitory I-units, which fire less when stimulated by inhibitory interneurons activated by P-units.
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Kawasaki, M. (1975). "Independently evolved jamming avoidance responses in Gymnotid and Gymnarchid electric fish: a case of convergent evolution of behavior and its sensory basis".
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are time coders, like the T-units in Gymnotiformes. O-units code the signal's intensity, like P-units in Gymnotiformes, but respond over a narrower range of intensities. In
1147:"Phase and amplitude computations in the midbrain of an electric fish: Intracellular studies of neurons participating in the jamming avoidance response of Eigenmannia" 415:, omitting non-electric and strongly-electric fishes, shows major events in their evolution. In the tree, "sp" means "a species" and "spp" means "multiple species". 181:– they can locate objects by generating an electric field and detecting distortions in the field caused by interference from those objects. Electric fish use their 1235:(1975). "Comparison of the jamming avoidance responses in Gymnotid and Gymnarchid electric fish: A case of convergent evolution of behavior and its sensory basis". 411:
nearly identical neural computational mechanisms and behavioral responses to avoid jamming, with only minor differences. The phylogeny of the weakly electric fish
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are very similar, each fish shifts its discharge frequency to increase the difference between the two. By doing this, both fish prevent jamming of their sense of
1444:(1977) Principles of Electrolocation and Jamming Avoidance in Electric Fish: A Neuroethological Approach. Studies of Brain Function, Vol. 1. Berlin-New York: 996:
Tan, E.; Nizar, J.; Carrera-G, E.; Fortune, E. (2005). "Electrosensory interference in naturally occurring aggregates of a species of weakly electric fish,
949:"The complexity of high-frequency electric fields degrades electrosensory inputs: implications for the jamming avoidance response in weakly electric fish" 1380:
Lavoué, Sébastien; Miya, Masaki; Arnegard, Matthew E.; Sullivan, John P.; Hopkins, Carl D.; Nishida, Mutsumi (2012-05-14). Murphy, William J. (ed.).
190:, frequencies range from 240 to 600 Hz. The EOD frequency is very steady, typically with less than 0.3% variation over a 10-minute time span. 104:
The jamming avoidance response (JAR) was discovered by Akira Watanabe and Kimihisa Takeda in 1963. The fish they used was an unspecified species of
338:, phase differences between EOD and stimulus are calculated in the electrosensory lateral line lobe rather than in the torus semicircularis. 1052:; Hamstra, R. Jr. (1973). "Coding properties of two classes of afferent nerve fibers: high-frequency electroreceptors in the electric fish, 263:, and the electroreceptive organs perceive a single wave with an intermediate frequency. In addition, the combined stimulus-EOD wave has a 2478: 1675: 2220: 2816: 1494: 199:
between their discharges. Therefore, it seems to be the function of the JAR to avoid sensory confusion among neighboring fish.
1277:"The phylogenetic distribution of electroreception: Evidence for convergent evolution of a primitive vertebrate sense modality" 1196:
Kramer, Berndt (15 May 1999). "Waveform discrimination, phase sensitivity and jamming avoidance in a wave-type electric fish".
2844: 2047: 1517: 1104: 871: 869:(1975). "Electrolocation and jamming avoidance in the electric fish Gymnarchus niloticus (Gymnarchidae, Mormyriformes)". 796: 768: 684: 2861: 2077: 1337: 1237: 831: 1382:"Comparable Ages for the Independent Origins of Electrogenesis in African and South American Weakly Electric Fishes" 259:
When the stimulus frequency and discharge frequency are close to each other, the two amplitude-time waves undergo
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which fires at the signal frequency, and a P-unit tuberous receptor which fires fastest near the peak of the
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to have its neural circuitry completely specified. As such, it holds special significance in the field of
2809: 1716: 1487: 1151: 829:; Hamstra, R. Jr.; Scheich, H. (1972). "The jamming avoidance response of high frequency electric fish". 788: 3065: 2468: 2195: 2165: 1661: 1276: 1058: 1002: 2866: 2741: 2734: 2701: 2443: 2115: 1796: 1632: 3070: 2751: 2510: 1538: 182: 2923: 2729: 2711: 2273: 2022: 1773: 1049: 826: 124: 21: 18:
Behavior performed by weakly electric fish to prevent jamming of their sense of electroreception
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is nearly identical to that of the Gymnotiformes, with a few minor differences. S-units in
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meet, one fish shifts its frequency upward and the other shifts its frequency downward.
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Most of the JAR pathway in the South American Gymnotiformes has been worked out using
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Hopkins, C. (1974). "Electric communication: functions in the social behavior of
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Kawasaki, M. (2009) Evolution of time-coding systems in weakly electric fishes.
551: 3075: 2948: 2943: 2891: 2668: 2641: 2533: 2525: 2458: 2428: 2367: 2349: 2303: 2293: 1897: 1850: 432: 229: 1102:(1980). "Neural correlates of the jamming avoidance response of Eigenmannia". 1015: 511: 3144: 2938: 2871: 2825: 2770: 2673: 2587: 2407: 2382: 2377: 2333: 2328: 2283: 2278: 2258: 2130: 1909: 1828: 1575: 1503: 763: 597: 378: 361: 357: 347: 299: 169: 93: 69: 59: 1209: 1071: 925: 789:"The change of discharge frequency by A.C. stimulus in a weak electric fish" 679: 3095: 3045: 2983: 2663: 2487: 2402: 2362: 2263: 2205: 2145: 2072: 2067: 2057: 1973: 1963: 1892: 1875: 1782: 1743: 1736: 1637: 1581: 1569: 1551: 1425: 1217: 1023: 982: 964: 657: 649: 619: 566: 534: 526: 469: 311: 118:. They found that when a sinusoidal electrical stimulus is emitted from an 31:
perform the jamming avoidance response: When two fish with around the same
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Sign-selective cells input into the nucleus electrosensorius in the
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caused by the interference of two signals with similar frequencies.
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Gymnotiforms have two classes of electroreceptive organs, the
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Bullock, T. H.; Bodznick, D. A.; Northcutt, R. G. (1983).
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experimentally by silencing a fish's electric organ with
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Spherical cells and pyramidal cells then project to the
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Heiligenberg, W. (1991) Neural Nets in Electric fish.
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The behavior has been most intensively studied in the
1132: 1047: 825: 1097: 1451:Heiligenberg, W. (1990) Electric Systems in Fish. 940: 3142: 1141: 947:Shifman, Aaron R.; Lewis, John E. (2018-01-04). 859: 298:, a structure with many laminae (layers) in the 342:Phylogeny and evolution of weakly electric fish 786: 2810: 1669: 1488: 946: 865: 1189: 395:in Apteronotoidea that perform JAR include 127:, Robert Hamstra Jr., and Henning Scheich. 2817: 2803: 1676: 1662: 1495: 1481: 787:Watanabe, Akira; Takeda, Kimihisa (1963). 281: 150:discovered a JAR in the distantly-related 2221:Tradeoffs for locomotion in air and water 1415: 1405: 1303: 1172: 972: 1334: 239: 129: 20: 905: 138:also has a jamming avoidance response, 3143: 1195: 953:Journal of the Royal Society Interface 286:The time-coding T-units converge onto 2798: 1657: 1476: 3123: 2777: 2048:Electroreception and electrogenesis 1518:Electroreception and electrogenesis 1105:Journal of Comparative Physiology A 872:Journal of Comparative Physiology A 797:The Journal of Experimental Biology 769:Electroreception and electrogenesis 177:and other weakly electric fish use 13: 1435: 1165:10.1523/JNEUROSCI.05-02-00515.1985 14: 3167: 2824: 1502: 1338:Journal of Comparative Physiology 1238:Journal of Comparative Physiology 832:Journal of Comparative Physiology 608:S. American knifefishes 235: 43:is a behavior of some species of 3122: 3111: 3110: 2776: 2765: 2764: 2747: 2746: 1772: 683: 678: 623: 618: 555: 550: 515: 510: 451: 446: 2867:Central pattern generator (CPG) 1732:Environmental impact of fishing 1328: 1224: 1198:Journal of Experimental Biology 210: 68:. It is also present in other 3061:Frog hearing and communication 989: 899: 780: 1: 774: 326:The neural pathway of JAR in 1817:intramembranous ossification 1407:10.1371/journal.pone.0036287 1296:10.1016/0165-0173(83)90003-6 114:wave discharge of about 300 99: 7: 1462:: Cambridge, Massachusetts. 1152:The Journal of Neuroscience 757: 163: 10: 3172: 3066:Infrared sensing in snakes 3051:Jamming avoidance response 2196:Fin and flipper locomotion 2166:Sequential hermaphroditism 2053:Jamming avoidance response 1770: 1578:(S. American knifefishes) 1523:Jamming avoidance response 1231:Bullock, T.; Behrend, K.; 1059:Journal of Neurophysiology 1003:Behavioural Brain Research 345: 167: 160:in two separate lineages. 41:jamming avoidance response 3106: 3033: 3012: 2916: 2832: 2760: 2684: 2617: 2524: 2486: 2477: 2416: 2347: 2234: 2186: 2096: 2021: 1781: 1691: 1633:History of bioelectricity 1625: 1562: 1531: 1510: 1016:10.1016/j.bbr.2005.06.014 671: 661: 611: 601: 591: 584: 538: 500: 490: 480: 473: 436: 426: 356:of weakly electric fish, 305: 3071:Caridoid escape reaction 1683: 2924:Theodore Holmes Bullock 2712:Glossary of ichthyology 2274:Diel vertical migration 1210:10.1242/jeb.202.10.1387 1072:10.1152/jn.1973.36.1.39 926:10.1163/156853974X00499 282:Processing in the brain 3081:Surface wave detection 2078:Surface wave detection 2043:Hydrodynamic reception 1717:Diseases and parasites 1284:Brain Research Reviews 965:10.1098/rsif.2017.0633 256: 179:active electrolocation 143: 36: 3151:Animal nervous system 2897:Anti-Hebbian learning 2216:Undulatory locomotion 2033:Ampullae of Lorenzini 1547:Ampullae of Lorenzini 998:Eigenmannia virescens 908:Eigenmannia virescens 668:Electric catfish 346:Further information: 243: 226:Eigenmannia virescens 168:Further information: 133: 65:Eigenmannia virescens 24: 2974:Bernhard Hassenstein 2907:Ultrasound avoidance 2882:Fixed action pattern 2845:Coincidence detector 2444:Genetically modified 1606:(electric catfishes) 1233:Heiligenberg, Walter 1143:Heiligenberg, Walter 1100:Heiligenberg, Walter 1050:Bullock, Theodore H. 867:Heiligenberg, Walter 827:Bullock, Theodore H. 441:(skates) (~200 spp) 409:convergently evolved 384:Gymnarchus niloticus 296:torus semicircularis 158:convergently evolved 153:Gymnarchus niloticus 140:evolved convergently 110:, which has a quasi- 85:Gymnarchus niloticus 78:, as well as in the 45:weakly electric fish 3041:Animal echolocation 2979:Werner E. Reichardt 2929:Walter Heiligenberg 2249:Aquatic respiration 2136:Life history theory 1572:(African knifefish) 1398:2012PLoSO...736287L 1145:; Rose, G. (1985). 810:10.1242/jeb.40.1.57 381:), and the species 352:There are two main 322:(Osteoglossiformes) 272:ampullary receptors 148:Walter Heiligenberg 3004:Fernando Nottebohm 2902:Sound localization 2877:Lateral inhibition 2088:Weberian apparatus 1467:Zoological Science 1351:10.1007/BF00209614 1251:10.1007/BF01380047 1118:10.1007/BF00656908 885:10.1007/bf01380044 845:10.1007/BF00696517 276:tuberous receptors 257: 144: 37: 3138: 3137: 3025:Slice preparation 2887:Krogh's Principle 2862:Feature detection 2792: 2791: 2702:Fish common names 2613: 2612: 2244:Aquatic predation 2068:Capacity for pain 1797:Age determination 1651: 1650: 1543:Electroreceptors 1204:(10): 1387–1398. 959:(138): 20170633. 754: 753: 745: 744: 732: 731: 723: 722: 714: 713: 705: 704: 696: 695: 636: 635: 573: 572: 542:African knifefish 487:Osteoglossiformes 389:Osteoglossiformes 366:Osteoglossiformes 249:tuberous receptor 136:African knifefish 3163: 3126: 3125: 3114: 3113: 3091:Mechanoreception 3086:Electroreception 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472: 466: 465: 458: 457: 435: 433:Chondrichthyes 427: 425: 417: 343: 340: 323: 316: 307: 304: 283: 280: 237: 236:Sensory coding 234: 221: 214: 212: 209: 183:electric organ 165: 162: 101: 98: 60:South American 17: 9: 6: 4: 3: 2: 3168: 3157: 3156:Neuroethology 3154: 3152: 3149: 3148: 3146: 3131: 3130: 3121: 3119: 3118: 3109: 3108: 3105: 3097: 3094: 3093: 3092: 3089: 3087: 3084: 3082: 3079: 3077: 3074: 3072: 3069: 3067: 3064: 3062: 3059: 3057: 3054: 3052: 3049: 3047: 3044: 3042: 3039: 3038: 3036: 3032: 3026: 3023: 3021: 3018: 3017: 3015: 3011: 3005: 3002: 3000: 2997: 2995: 2992: 2990: 2987: 2985: 2982: 2980: 2977: 2975: 2972: 2970: 2967: 2965: 2962: 2960: 2957: 2955: 2952: 2950: 2947: 2945: 2942: 2940: 2939:Konrad Lorenz 2937: 2935: 2932: 2930: 2927: 2925: 2922: 2921: 2919: 2915: 2908: 2905: 2903: 2900: 2898: 2895: 2893: 2890: 2888: 2885: 2883: 2880: 2878: 2875: 2873: 2872:NMDA receptor 2870: 2868: 2865: 2863: 2860: 2858: 2855: 2853: 2852: 2848: 2846: 2843: 2841: 2838: 2837: 2835: 2831: 2827: 2826:Neuroethology 2820: 2815: 2813: 2808: 2806: 2801: 2800: 2797: 2785: 2784: 2775: 2773: 2772: 2763: 2762: 2759: 2753: 2752:more lists... 2745: 2743: 2740: 2736: 2733: 2731: 2728: 2727: 2725: 2723: 2720: 2718: 2715: 2713: 2710: 2708: 2707:Fish families 2705: 2703: 2700: 2698: 2695: 2693: 2692:Aquarium life 2690: 2689: 2687: 2683: 2675: 2674:fleshy-finned 2672: 2670: 2667: 2666: 2665: 2662: 2658: 2655: 2653: 2650: 2648: 2645: 2644: 2643: 2642:Cartilaginous 2640: 2636: 2633: 2631: 2628: 2627: 2626: 2623: 2622: 2620: 2616: 2604: 2601: 2599: 2596: 2594: 2591: 2590: 2589: 2586: 2582: 2579: 2577: 2574: 2572: 2569: 2567: 2564: 2563: 2562: 2559: 2555: 2552: 2550: 2547: 2545: 2542: 2540: 2537: 2536: 2535: 2532: 2531: 2529: 2527: 2523: 2517: 2514: 2512: 2509: 2507: 2504: 2502: 2499: 2497: 2494: 2493: 2491: 2489: 2485: 2482: 2480: 2476: 2470: 2467: 2465: 2462: 2460: 2457: 2455: 2452: 2450: 2447: 2445: 2442: 2440: 2437: 2435: 2432: 2430: 2427: 2425: 2422: 2421: 2419: 2415: 2409: 2406: 2404: 2401: 2399: 2396: 2394: 2391: 2389: 2386: 2384: 2381: 2379: 2376: 2374: 2371: 2369: 2366: 2364: 2361: 2359: 2356: 2355: 2353: 2351: 2346: 2340: 2337: 2335: 2332: 2330: 2327: 2325: 2322: 2320: 2317: 2315: 2312: 2310: 2307: 2305: 2302: 2300: 2297: 2295: 2292: 2290: 2287: 2285: 2282: 2280: 2279:Electric fish 2277: 2275: 2272: 2270: 2267: 2265: 2262: 2260: 2257: 2255: 2252: 2250: 2247: 2245: 2242: 2241: 2239: 2233: 2227: 2224: 2222: 2219: 2217: 2214: 2212: 2209: 2207: 2204: 2202: 2199: 2197: 2194: 2193: 2191: 2189: 2185: 2177: 2174: 2173: 2172: 2169: 2167: 2164: 2162: 2159: 2157: 2154: 2152: 2149: 2147: 2144: 2142: 2139: 2137: 2134: 2132: 2129: 2127: 2124: 2122: 2119: 2117: 2114: 2112: 2109: 2107: 2104: 2103: 2101: 2099: 2095: 2089: 2086: 2084: 2081: 2079: 2076: 2074: 2071: 2069: 2066: 2064: 2061: 2059: 2056: 2054: 2051: 2049: 2046: 2044: 2041: 2039: 2036: 2034: 2031: 2030: 2028: 2026: 2020: 2014: 2011: 2009: 2006: 2002: 1999: 1997: 1994: 1993: 1992: 1989: 1985: 1982: 1980: 1977: 1976: 1975: 1972: 1970: 1967: 1965: 1962: 1958: 1955: 1954: 1953: 1950: 1948: 1945: 1943: 1940: 1938: 1935: 1933: 1930: 1926: 1923: 1922: 1921: 1918: 1916: 1913: 1911: 1910:Mauthner cell 1908: 1906: 1903: 1899: 1896: 1894: 1891: 1890: 1889: 1886: 1884: 1881: 1877: 1874: 1872: 1869: 1867: 1864: 1862: 1859: 1857: 1854: 1852: 1849: 1848: 1847: 1844: 1840: 1837: 1836: 1835: 1832: 1830: 1829:Chromatophore 1827: 1825: 1822: 1818: 1815: 1813: 1810: 1809: 1808: 1805: 1803: 1800: 1798: 1795: 1794: 1792: 1790: 1784: 1780: 1775: 1765: 1762: 1760: 1757: 1755: 1752: 1750: 1747: 1745: 1742: 1738: 1735: 1733: 1730: 1728: 1725: 1724: 1723: 1720: 1718: 1715: 1713: 1710: 1708: 1705: 1703: 1700: 1699: 1697: 1695: 1690: 1686: 1679: 1674: 1672: 1667: 1665: 1660: 1659: 1656: 1644: 1641: 1639: 1636: 1634: 1631: 1630: 1628: 1624: 1617: 1616:Uranoscopidae 1614: 1611: 1608: 1605: 1602: 1599: 1596: 1593: 1590: 1585: 1584: 1583:Electrophorus 1580: 1579: 1577: 1576:Gymnotiformes 1574: 1571: 1568: 1567: 1565: 1561: 1553: 1550: 1548: 1545: 1544: 1542: 1540: 1537: 1536: 1534: 1530: 1524: 1521: 1519: 1516: 1515: 1513: 1509: 1505: 1504:Electric fish 1498: 1493: 1491: 1486: 1484: 1479: 1478: 1475: 1468: 1464: 1461: 1457: 1454: 1450: 1447: 1443: 1440: 1439: 1427: 1423: 1418: 1413: 1408: 1403: 1399: 1395: 1392:(5): e36287. 1391: 1387: 1383: 1376: 1368: 1364: 1360: 1356: 1352: 1348: 1345:(1): 97–121. 1344: 1340: 1339: 1331: 1323: 1319: 1315: 1311: 1306: 1305:2027.42/25137 1301: 1297: 1293: 1289: 1285: 1278: 1271: 1269: 1260: 1256: 1252: 1248: 1244: 1240: 1239: 1234: 1227: 1219: 1215: 1211: 1207: 1203: 1199: 1192: 1184: 1180: 1175: 1170: 1166: 1162: 1158: 1154: 1153: 1148: 1144: 1138: 1136: 1127: 1123: 1119: 1115: 1111: 1107: 1106: 1101: 1098:Bastian, J.; 1094: 1092: 1090: 1081: 1077: 1073: 1069: 1065: 1061: 1060: 1055: 1051: 1048:Scheich, H.; 1044: 1042: 1033: 1029: 1025: 1021: 1017: 1013: 1009: 1005: 1004: 999: 992: 984: 980: 975: 970: 966: 962: 958: 954: 950: 943: 935: 931: 927: 923: 919: 915: 914: 909: 902: 894: 890: 886: 882: 878: 874: 873: 868: 862: 854: 850: 846: 842: 838: 834: 833: 828: 822: 820: 811: 807: 803: 799: 798: 790: 783: 779: 770: 767: 765: 764:Electric fish 762: 761: 750: 749: 741: 737: 736: 728: 727: 719: 718: 710: 709: 701: 700: 692: 691: 688: 686: 681: 676: 669: 666: 665: 659: 656: 655: 652: 651: 646: 645: 641: 640: 632: 631: 628: 626: 621: 616: 609: 606: 605: 599: 598:Gymnotiformes 596: 595: 589: 588: 582: 581: 578: 577: 569: 568: 567:knollenorgans 564: 563: 560: 558: 553: 548: 544: 543: 536: 533: 532: 529: 528: 527:knollenorgans 524: 523: 520: 518: 513: 508: 504: 498: 495: 494: 488: 485: 484: 478: 477: 471: 468: 467: 464: 460: 459: 456: 454: 449: 444: 440: 434: 431: 430: 424: 421: 420: 416: 414: 410: 406: 405: 400: 399: 394: 390: 386: 385: 380: 379:Gymnotiformes 375: 371: 367: 363: 362:South America 359: 358:Gymnotiformes 355: 349: 348:Electric fish 339: 337: 333: 329: 321: 315: 313: 303: 301: 300:mesencephalon 297: 292: 289: 279: 277: 273: 268: 266: 262: 254: 250: 246: 242: 233: 231: 227: 219: 208: 206: 200: 197: 191: 189: 184: 180: 176: 171: 170:Electric fish 161: 159: 155: 154: 149: 141: 137: 132: 128: 126: 121: 117: 113: 109: 108: 97: 95: 94:neuroethology 91: 87: 86: 81: 77: 76: 71: 70:Gymnotiformes 67: 66: 61: 56: 54: 50: 46: 42: 34: 30: 29: 23: 16: 3127: 3115: 3096:Lateral line 3050: 3046:Waggle dance 2984:Eric Knudsen 2849: 2781: 2769: 2669:spiny-finned 2618:Major groups 2339:Intelligence 2319:Scale eaters 2264:Cleaner fish 2146:Mouthbrooder 2098:Reproduction 2073:Schreckstoff 2058:Lateral line 2052: 1974:Swim bladder 1964:Spiral valve 1893:hyomandibula 1876:pseudobranch 1759:Hypoxia in - 1638:Lateral line 1618:(stargazers) 1582: 1570:Gymnarchidae 1552:Knollenorgan 1522: 1469:26: 587-599. 1389: 1385: 1375: 1342: 1336: 1330: 1290:(1): 25–46. 1287: 1283: 1242: 1236: 1226: 1201: 1197: 1191: 1156: 1150: 1109: 1103: 1066:(1): 39–60. 1063: 1057: 1053: 1007: 1001: 997: 991: 956: 952: 942: 917: 911: 907: 901: 879:(1): 55–67. 876: 870: 861: 836: 830: 801: 795: 782: 674: 672: 658:Siluriformes 648: 642: 614: 612: 607: 565: 546: 540: 539: 535:Gymnarchidae 525: 506: 501: 442: 437: 402: 396: 391:). Notable 382: 351: 335: 331: 327: 325: 319: 312:diencephalon 309: 293: 285: 269: 265:beat pattern 261:interference 258: 244: 230:model system 225: 223: 217: 211:Neurobiology 201: 195: 192: 187: 174: 173: 151: 145: 105: 103: 83: 73: 63: 57: 40: 38: 26: 15: 3020:Patch clamp 2989:Eric Kandel 2969:Franz Huber 2840:Feedforward 2783:WikiProject 2742:Prehistoric 2726:Threatened 2417:Other types 2314:Sardine run 2289:Forage fish 2269:Corallivory 2121:Development 2106:Bubble nest 1979:physoclisti 1969:Suckermouth 1942:Root effect 1764:Ichthyology 1054:Eigenmannia 839:(1): 1–22. 505:(~200 spp) 423:Vertebrates 404:Apteronotus 398:Eigenmannia 318:Pathway in 245:Eigenmannia 218:Eigenmannia 216:Pathway in 196:Eigenmannia 188:Eigenmannia 175:Eigenmannia 107:Eigenmannia 75:Apteronotus 49:frequencies 28:Eigenmannia 3145:Categories 2994:Nobuo Suga 2909:in insects 2398:Groundfish 2393:Freshwater 2388:Euryhaline 2373:Coral reef 2309:Salmon run 2299:Paedophagy 2201:Amphibious 2188:Locomotion 1996:pharyngeal 1984:physostome 1937:Photophore 1883:Glossohyal 1856:gill raker 1839:dorsal fin 1789:physiology 1598:Rajiformes 1592:Mormyridae 1511:Physiology 1455:6:196-206. 775:References 497:Mormyridae 439:Rajiformes 336:Gymnarchus 332:Gymnarchus 328:Gymnarchus 320:Gymnarchus 253:beat cycle 112:sinusoidal 90:vertebrate 2647:chimaeras 2534:Predatory 2511:Salmonids 2469:Whitefish 2459:Poisonous 2434:Diversity 2368:Coldwater 2304:Predatory 2294:Migratory 2254:Bait ball 2237:behaviour 2156:Pregnancy 2151:Polyandry 1925:papillare 1920:Operculum 1915:Meristics 1861:gill slit 1824:Cleithrum 1754:Fish kill 1744:Fear of - 1737:- as food 1727:Fisheries 1712:Evolution 1702:Diversity 1460:MIT Press 913:Behaviour 804:: 57–66. 738:425  673:(11 spp) 644:ampullary 461:430  146:In 1975, 120:electrode 100:Discovery 33:frequency 3117:Category 2857:Instinct 2833:Concepts 2771:Category 2722:Smallest 2635:lampreys 2598:flatfish 2588:Demersal 2544:mackerel 2539:billfish 2479:Commerce 2408:Tropical 2383:Demersal 2378:Deep-sea 2334:Venomous 2226:RoboTuna 2176:triggers 2171:Spawning 2131:Juvenile 2116:Egg case 1749:FishBase 1600:(skates) 1426:22606250 1386:PLOS ONE 1367:27622365 1322:15603518 1259:20094087 1218:10210679 1126:26069930 1032:15744149 1024:16099058 983:29367237 893:42982465 853:19690508 758:See also 650:tuberous 274:and the 164:Behavior 82:species 72:such as 62:species 3129:Commons 3034:Systems 3013:Methods 2717:Largest 2630:hagfish 2625:Jawless 2603:pollock 2576:sardine 2571:herring 2566:anchovy 2516:Tilapia 2506:Octopus 2501:Catfish 2488:Farming 2403:Pelagic 2363:Coastal 2350:habitat 2206:Walking 2111:Clasper 2063:Otolith 2025:systems 2023:Sensory 1957:ganoine 1932:Papilla 1783:Anatomy 1722:Fishing 1626:Related 1532:Anatomy 1453:Synapse 1417:3351409 1394:Bibcode 1359:8366474 1314:6616267 1183:3973680 1174:6565201 1080:4705666 974:5805966 934:4533613 647:,  545:(1 sp) 387:(order 288:neurons 80:African 2917:People 2851:Umwelt 2735:sharks 2652:sharks 2581:sprats 2561:Forage 2549:salmon 2429:Coarse 2211:Flying 2083:Vision 2038:Barbel 1952:Scales 1812:dermal 1692:About 1563:Groups 1424:  1414:  1365:  1357:  1320:  1312:  1257:  1216:  1181:  1171:  1124:  1078:  1030:  1022:  981:  971:  932:  891:  851:  413:clades 393:genera 370:Africa 354:orders 306:Output 205:curare 2697:Blind 2685:Lists 2464:Rough 2329:Sleep 2235:Other 2001:shark 1991:Teeth 1363:S2CID 1318:S2CID 1280:(PDF) 1255:S2CID 1122:S2CID 1028:S2CID 930:JSTOR 889:S2CID 849:S2CID 792:(PDF) 675:Pulse 507:Pulse 443:Pulse 368:from 360:from 228:as a 2730:rays 2664:Bony 2657:rays 2554:tuna 2496:Carp 2454:Oily 2439:Game 2424:Bait 2358:Cave 2141:Milt 1846:Gill 1834:Fins 1807:Bone 1694:fish 1685:Fish 1422:PMID 1355:PMID 1310:PMID 1214:PMID 1179:PMID 1076:PMID 1020:PMID 979:PMID 615:Wave 547:Wave 401:and 364:and 134:The 39:The 2593:cod 2348:By 2161:Roe 1888:Jaw 1786:and 1412:PMC 1402:doi 1347:doi 1343:103 1300:hdl 1292:doi 1247:doi 1243:103 1206:doi 1202:202 1169:PMC 1161:doi 1114:doi 1110:136 1068:doi 1056:". 1012:doi 1008:164 1000:". 969:PMC 961:doi 922:doi 910:". 881:doi 877:103 841:doi 806:doi 740:mya 463:mya 372:. 3147:: 1420:. 1410:. 1400:. 1388:. 1384:. 1361:. 1353:. 1341:. 1316:. 1308:. 1298:. 1286:. 1282:. 1267:^ 1253:. 1241:. 1212:. 1200:. 1177:. 1167:. 1155:. 1149:. 1134:^ 1120:. 1108:. 1088:^ 1074:. 1064:36 1062:. 1040:^ 1026:. 1018:. 1006:. 977:. 967:. 957:15 955:. 951:. 928:. 918:50 916:. 887:. 875:. 847:. 837:77 835:. 818:^ 802:40 800:. 794:. 677:: 617:: 549:: 509:: 445:: 232:. 116:Hz 96:. 55:. 2818:e 2811:t 2804:v 1677:e 1670:t 1663:v 1496:e 1489:t 1482:v 1448:. 1428:. 1404:: 1396:: 1390:7 1369:. 1349:: 1324:. 1302:: 1294:: 1288:6 1261:. 1249:: 1220:. 1208:: 1185:. 1163:: 1157:5 1128:. 1116:: 1082:. 1070:: 1034:. 1014:: 985:. 963:: 936:. 924:: 895:. 883:: 855:. 843:: 812:. 808:: 142:.

Index

diagram showing change in waves emitted by two electric fish to avoid each other's frequency
Eigenmannia
frequency
weakly electric fish
frequencies
electroreception
South American
Eigenmannia virescens
Gymnotiformes
Apteronotus
African
Gymnarchus niloticus
vertebrate
neuroethology
Eigenmannia
sinusoidal
Hz
electrode
Theodore Bullock
photo of an electric fish
African knifefish
evolved convergently
Walter Heiligenberg
Gymnarchus niloticus
convergently evolved
Electric fish
active electrolocation
electric organ
curare
model system

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