4 corp, Physiol. 128, 199- 192 (1978) Key Stimuli of Prey for Toads (Bufo bufo L.) Configuration and Movement Patterns* H.-W. Borchers, H. Burghagen, and J.-P. Ewert Neurvethology anu Blocybemetie Laboratories, University of Kassel, D390 Kussel, Federal Republic of Germany Accepued! August 22, 1978 Summary. The ability of the Common Toad Bufo Bufo (L.} to distinguish between a worm-like and an antiworm-like (e.g,, a bar at right angles to its direc- tion of motion) moving stripe is not generally altered by movement dynamics, such as velocity of motion or by particular movement patterns—so far as has been investigated. A small square stimulus, however, with its “indifferent” shape is more attractive as prey for the toad when displaced in a stepwise manner, rather than moved at corresponding continuous (con- stant) velocity. Step frequencies of | 10 2eps were found to have optimal releasing values. Introduction Toads Bufo bufo (L.) distinguish visual prey objects from predators or behaviorally irrelevant objects on the basis of configurational stimulus parameters linked with the direction of stimulus movement (for review sce Evert, 1976), Our previous experiments have been performed with stimuli moved contin- uously at constunt velocities. Considering the richness of movement patterns of the natural prey objects, however, our experimental stinmulus movements seem. to represent only @ crude simplification. Sensitivity to jerky movements was recenily observed in the ¥i- sually guided prey-catching behavior of salamanders (Roth, 1978). The present study with Common Toads Bufo bufo (L.) investigates the influence of movement dynamics on the discrimination of moving configura- téonai stimuli. Method Stinutas Configurations The standard stimuli are shown in Fig. 2A CA black 3330 ramn? stripe moved in 2 worWhke Way Wid is long ax pave) te + Supported by the Deutiche Forschungssemeinschall Fav 7/6 the Horizontal dircstion of motion (A), & black 329 mm? stipe moved antisoraike sith sts axis perpendicular to the ditcotion fof moiion (B). and a black GGmm? square (C1. AU stimuli ‘Were moved on 4 white background at stance of 7 ex, [rom ‘the toad’s eyes. The amount af the stimulus background coutrst wae C=Ly L/L, L409 with £ led xm 2) businances of the stimulus (3) nd the buckground (9), Movement Patteras ‘The stimulus was moved linearly in «horizontal (1-) direction by a motor-driven belt system (Fig. 1. Movement patterns so called step_movements—nere generited by different waveform Tunetions (Fag. 23-0): Square wave (a, triangular (5) or sinaxendal fe). The pulses of the waveforms copttollad the velocity of the bell-moter (FD. The ampicudes of the pukes corresponded ley the stmmulus movement veloeiy. fase baseline Bee) a the plkes wus vero (Fig. 2 abuvel, The maximum pest-lo-peak devel was the samme in all exses {Fig 2a-c) and it corresponded to Mims movement velocity {rq} 9¢ 8 mans, Thus stinoull eel bhemovedin steps by 3 different mods Tower races in Fig 23 ©) Fig. 1. Faperimenial prosslute for measrements of (he tds (Bufo hugo) prey-catching vccisey in response £0 a blick stiesuhss object (3) moved in hovizoaual (x) digestion feom left wo right ‘on a white belt (O), The belt is driven continuousty hy aa cietronic ddevies consisting of an clecirc motar (M). a tachogencrator (TG) and « controller Ci, Stepemovements can be geacrated By means ‘of funtion generator (1) which gives the reference inpa for ‘the covtroller (0340-7594/78/0128/0189/S01,00HLAW, Boreners etal : Configuration and Movement Patterns fal {h) {c} 3 Pulse Amplitude Colts has Movement Velocity Cmm/s} ? 3 a 8 zg Zio aj = 23 in 2 2 Ol ome! | onoenonm—n! c = | inn ‘ IN a a) Mm mor tm moot 1 mt ‘Step -movement Frequency (cps| Fig. 2. tnilucnse of stimulas objcts of dierent confignritions (4 C) moved in stepwise manner hy diferent sep froquencis ane functions (a «) on the prey-caleking neiening activity ofthe toa Bufo bia. Cach curve point represents average response activities from 2slfercat animals Upper races: Relerence ie of the controler ésez Fig. 1). Lower tees: Quipes valkage of the sechogenerator ay sav inden of the movement sslock rg, of the stimulls. (The pple 30 the revards 1s past of the pulsating DC voltae of the ihagsrerator. Dislrlin at 1H) ops may be aepested In the present experiments becwuse dh iss visual system is able te eesse these frequency pattern) Records sbowe: Slep moveneol frequencies of 2eps: records below: Step movement frequencies of 3

The averane Hep movement velocity ws constant at £,—9 mms in all cases, Mocement Dynamics Fig. 2a 6) At coustant frequency, in (a} the stimulus velocity within a hall fyele remained constant. in (b) the velocity within & balf eyele ‘aried with constant acceleration, and in (e) the neeeleration within the eycle varied cosinusoidaly, By increasing the frequency in {a the stimulus velocity within « half cyule remained constant, (6) the amount of the acceleration increased and 5 (e) the chasse fof tie acceleration imereased, Measure of the Releasing Value of the Stimulus During the experiment the oad sit within a cylindrical glass vessel in front of the helt systoo (Fig. 1) he stimu ited the category prey he toad responded with orienting movements of the head and body toward the stimulus object. The number of prev-catching orienting movenicnts per traverse of the stimulus through 2 froaral pat of the binocular visual field of (04 visual angle (-280 mm, ‘and 4 time ierval oF 2 8) was taken asi» measure of the Felsusing vas of the stimulus as a proy objeet Fepeciments were performed with a tolal of 20 roads. The influence of moverneat functions was tested in three groups ‘Fig, 24 c) being sepurated by 24 recovery periods, Within each group the frequency ofthe wavclorm Functions was chiaged saoce Svely in random erdor Results Responses to the wormlike object are shown in Fig. 2A, a-c, The average prey-catching activity of the toads was increased slightly when square (2), triangle (b) or sinusoidal generated step movements had frequencies in the range of 1-2 eps, The response activity to the antiworm (Fig, 2B, 4c) was zero for alt movement modes and step frequencies tested. In evaluating the results shown in Fig. 24 a possi- ble * suturation effect” should be considered: because the worm-like configuration is such a strong prey stimulus for the toad it iy not possible to increase the catching activity any further. Therefore compar tive experiments with a worm stimulus of same size but weak contrast (C*0.5) were carried out. In this case the level of prey-catching activity was decreased, but the effets of step movements showed no remark- able difference rom the previous results. In another set of experiments the worm and anti- worm stimuli were moved at constant (continuous) velocity tye=4.5, 9, 18 or 36 mm/s. The toads showed a clear preference for the worm configuration. The probability that the toads made a “misinterpre- Vion” —tracking an antiworm—in the present sclec- tion test at all movement velocities investigated was p< 0.01 (Hest) 1 The prey-catching activity in response to the square (Fig. 2C, a-c) averaged less than was obtained for the wormlike object. Surprisingly, here the prey- catching activity showed x statistically significam de- pendence on the step-movement frequency. For all Guce movement modes (4-c), step frequencies of 1-2 ops were found to be optimal rcleascrs (p< 0.0L; Hest). Step-movernents of 190 cps had the same weak releasing value as stimuli moved continuously at 9inmjs. These step frequencies cannot be resolved by the visual system of the toad. The value is far above threshold of the fusion frequency (Ewert, 196% Griisser-Cornehls, 1968). Further experiments with the square object were carried oul to answer the question of whether the tirme- pattern of the particular spatiat displacement pattern ofthe stimulus was the significant feature. The results showed that in the investigated velocity range 45<0,36 mm/s of square wave pulsc-generated Step movement patterns the optimal frequency of 1 2eps did not alter in the higher or lower ranges by a statistically significant amount, This indicates thal the time pattern rather than the spatial displace- ment pattern of the steps is the important stimulus feature, Discussion The results show that the ability of the toad Bufo ufo to distinguish worm-like from antiworm-like ob- jects is not altered by the dynamics of the stimulus— angular velocity and displacement pattern so far as has been investigated. In case of an “indillerent prey stimulus” configuration, such as with a square, the movement pattern may have strong effects on the releasing value. Thus, prey selection by the toad Bufo puso may depend on both configuration and move- ment pattern, J.¥, Lettvin writes: “frogs will leap to capture any object the size of an insect or worm, providing it moves like one” (Lettvin et al, 1959). Ewert and Kallweidt (in preparation) recently found in the frog Rana temporaria (L.) that particular move- snent patterns of a visual contrast stimulus. such as step movements (I step/s), are important features for ‘a stimulus to be categorized as prey. Similar results were obtained by D. Ingle (personal communication, 1977). Roth, recently working in our laboratory, observed also an influence of jerky movements on Ghe prey catching activity in salamanders Mydro- mantes genei (Roth, 1978). The movement paiterns investigated in all these studies, however, are prestm- ably only a part of a whole complex. In our future rescarch the movement dynamics of natural prey ob- jects must be analyzed and possibly compared with the optimal step frequencies of 1-2 cps obtained in