Journal of Comparative Psychology Manuscript version of Like Chimpanzees (Pan troglodytes), Pigeons (Columba livia domestica) Match and Nash Equilibrate Where Humans (Homo sapiens) Do Not Yosuke Hachiga, Lindsay P. Schwartz, Christopher Tripoli, Samuel Michaels, David Kearns, Alan Silberberg Funded by: • Japan Society • National Institutes of Health © 2018, American Psychological Association. This manuscript is not the copy of record and may not exactly replicate the final, authoritative version of the article. Please do not copy or cite without authors’ permission. The final version of record is available via its DOI: https://dx.doi.org/10.1037/com0000144 This article is intended solely for the personal use of the individual user and is not to be disseminated broadly. 1 1 2 Like Chimpanzees (Pan troglodytes), Pigeons (Columba livia domestica) Match and Nash Equilibrate Where Humans (Homo sapiens) Do Not 3 4 Yosuke Hachiga, Lindsay P. Schwartz, Christopher Tripoli, 5 Samuel Michaels, David Kearns, and Alan Silberberg 6 American University 7 8 9 10 11 12 13 14 15 16 Author Note Yosuke Hachiga, Lindsay P. Schwartz, Christopher Tripoli, Samuel Michaels, David Kearns, and Alan Silberberg, Department of Psychology, American University Correspondence concerning this article should be addressed to Yosuke Hachiga, Department of Psychology, American University, Washington, DC 20016. 17 Contact: hachi@american.edu 18 This work was funded by NIH 1 R15 MH109922 to American University and by JSPS 19 20 21 22 23 Overseas Research Fellowships to Yosuke Hachiga. 2 24 Abstract 25 Martin, Bhui, Bossaerts, Matsuzawa and Camerer (2014) found that chimpanzee pairs competing 26 in matching-pennies games achieved the Nash Equilibrium whereas human pairs did not. They 27 hypothesized this outcome may be due to: (a) chimpanzee ecology producing evolutionary 28 changes that give them a cognitive advantage over humans in these games; and (b) humans being 29 disadvantaged because the cognition necessary for optimal game play was traded off in evolution 30 to support language. We provide data relevant to their hypotheses by exposing pairs of pigeons 31 to the same games. Pigeons also achieved the Nash Equilibrium, but did so while also 32 conforming with the Matching-Law prediction on concurrent schedules that choice ratios covary 33 with reinforcer ratios. The cumulative-effects model, which produces matching on concurrent 34 schedules, also achieved the Nash Equilibrium when it was simulated on matching-pennies 35 games. The empirical and simulated compatibility between Matching-Law and Nash- 36 Equilibrium predictions can be explained in two ways. Choice to concurrent schedules, where 37 matching obtains, and choice in game play, where the Nash Equilibrium is achieved, may reflect 38 the operation of a common process in choice (e.g., reinforcer maximization) for which matching 39 and achieving the Nash Equilibrium are derivative. Alternatively, if matching in choice is innate 40 as some accounts argue, then achieving the Nash Equilibrium may be an epiphenomenon of 41 matching. Regardless, the wide species generality of matching relations in nonhuman choice 42 suggests game play in chimpanzees would not prove advantaged relative to most species in the 43 animal kingdom. 44 45 46 Keywords: Nash Equilibrium, matching pennies, matching law, chimpanzees, humans, pigeons 3 47 48 Like Chimpanzees (Pan troglodytes), Pigeons (Columba livia domestica) Match and Nash Equilibrate Where Humans (Homo sapiens) Do Not 49 Dennett (1983) has suggested that theorizing about mental-function levels in humans and 50 other animals is done by two types of researchers, killjoys and romantics. Killjoys place humans 51 at the apex of any ranking of mental function, viewing their mental capacities as unrivalled in the 52 animal kingdom; romantics, on the other hand, often see equivalences in mental function 53 between humans and some other animals. 54 We have no quarrel with this dichotomy and see the literature on comparative cognition 55 as offering cases where one or the other view seems sensible and useful. However, there is a 56 small literature that falls outside the Dennett dichotomy. This literature claims to show mental 57 function in chimpanzees that is superior to that seen in humans in terms of short-term memory 58 (Inoue & Matsuzawa, 2007), self-control in intertemporal preferences (Rosati, Stevens, Hare, & 59 Hauser, 2007) and rational maximizing (Jensen, Call, & Tomasello, 2007). 60 We have opposed these claims, arguing that, in one way or another, they overreach 61 (Genty, Karpel, & Silberberg, 2012; Silberberg & Kearns, 2009; Smith & Silberberg 2010). 62 These references once challenged all published claims of mental function outside the Dennett 63 dichotomy of which we were aware; however, matters have recently changed. Martin, Bhui, 64 Bossaerts, Matsuzawa, and Camerer (2014) offer credible demonstrations that when pairs of 65 chimpanzees or humans make repeated binary choices in competition for reinforcement, 66 chimpanzees come closer to achieving the so-called Nash Equilibrium (NE)—a point where each 67 participant in a competitive game cannot gain by a unilateral change in choice strategy if the 68 strategy of the opponent remains unchanged. To familiarize the reader with this work, we 69 summarize their report below. 4 70 Martin et al. (2014) procedure and results 71 Martin et al. (2014) exposed pairs of chimpanzees, sitting in proximity, to three 72 matching-pennies games in which each subject chose on its own touch screen between 73 illuminated left- and right-side squares. In all games, one member of the pair, the matcher, was 74 rewarded with apple only when it chose the same square on its touch screen as the other member; 75 the mismatcher, on the other hand, was rewarded only when it chose the square opposite that 76 chosen by the matcher. 77 The payoff structure differed in each game for the matcher: In the 2,000-trial symmetric 78 game, matches resulted in a single apple cube; in the 1,000-trial asymmetric game that followed, 79 left-side matches yielded three apple pieces and right-side matches yielded one; and finally, in 80 the 800-trial inspection game, left-side matches produced four apple pieces vs. one apple piece 81 for right-side matches. For mismatchers in these games, correct mismatches yielded one apple 82 piece in the symmetric game and two apple pieces in the other two games regardless of the side 83 chosen. 84 Martin et al. (2014) also exposed pairs of Japanese and African humans to the inspection 85 game for 200 trials as a matcher and 200 trials as a mismatcher, replacing food with money 86 amounts. The Japanese responded to the touch screens used with chimpanzees whereas the 87 Africans chose between bottle tops that were facing up or down. When bottle tops or side 88 preferences matched, the matcher was paid; when not, the mismatcher was paid. Payoffs on left 89 and right sides or up and down bottle tops mirrored those of the chimpanzees described above for 90 the inspection game. 91 Martin et al. (2014) calculated for each game the relative frequency of right-side or top- 92 up choices for chimpanzees and humans and compared it to the equilibrium point where neither 5 93 competitor could gain by a change in strategy if the competitor’s strategy remained unchanged. 94 This point, the NE, was approximated by the across-trials average performance of each 95 chimpanzee pair; however, the mismatching humans playing the inspection game chose the right 96 side or top up roughly 60% of the time, a value well short of the NE prediction of 80%. 97 Martin et al. (2014) speculated that the chimpanzee-human performance difference they 98 obtained was due to two factors. Regarding humans, evolutionary specializations such as 99 language and categorization skills required that other cognitive capacities, such as those used in 100 playing the inspection game, be compromised. They called this notion the “cognitive-tradeoff 101 hypothesis”, and it may account for why humans failed to achieve the NE on this task. 102 Regarding chimpanzees, their dominance-mediated social environment may have created a 103 fitness value for strength and speed in chimpanzees that translates into “a possible parallel 104 cognitive advantage… (p. 4)”. This added fitness value also may have aided chimpanzee 105 performance. 106 As noted above, our goal is to test psychological accounts that fall outside the Dennett 107 (1983) dichotomy. Martin et al.’s (2014) theorizing about the causes of chimpanzee superiority 108 in matching pennies certainly qualifies. Unfortunately, we see no way to test these accounts 109 individually: Are chimpanzees better at matching pennies than humans because humans have 110 been dumbed down by trading off task-specific cognitive function for language? Or are 111 chimpanzees smarter than humans because chimpanzees have a task-specific parallel cognitive 112 advantage? Either hypothesis rationalizes chimpanzee outperformance of humans. 113 Nevertheless, we do see a way these hypotheses can be tested jointly. The key is to choose a 114 species to match pennies so cognitively inferior to humans that readers will doubt its superior 6 115 performance, assuming it obtains, could plausibly be explained by either of Martin et al.’s 116 accounts. 117 Toward this end, we have selected pigeons to match pennies. Our argument is that even 118 if human mental facility specific to matching pennies has been compromised by the advent of 119 language, the consequences of that compromise could not credibly be so severe as to rationalize 120 why pigeons’ performance is superior to humans’. If pigeons are superior to humans in 121 matching pennies, this conceptual damage also extends to the notion of chimpanzees’ parallel 122 cognitive advantage because theorists that could believe chimpanzees have a task-specific 123 cognitive advantage would be less likely to extend this claim to pigeons. 124 This report compares chimpanzee approximations to the NE from Martin et al. (2014) 125 with those from pigeons on procedures like the ones they used with chimpanzees. As it happens, 126 pigeons have already been tested on a matching-pennies task by Sanabria and Thrailkill (2009). 127 They played a variety of matching-pennies games, but only two were also played in Martin et al. 128 (2014), the symmetric and asymmetric game. Based on the last session of performance in each 129 game from Sanabria and Thrailkill, pigeons appear to have matched chimpanzees in 130 approximating the NE in the symmetric game, but not in the asymmetric game. 131 Obviously, these data suggest that challenging Martin et al.’s (2014) two-theory account 132 of task-specific cognitive superiority in chimpanzees by showing pigeons are also superior to 133 humans may be empirically ill considered. However, we persist because we speculate that 134 pigeons’ failure to approximate the NE as well as chimpanzees may have been due to Sanabria 135 and Thrailkill’s (2009) procedure differing in a critical way from Martin et al.’s: Sanabria and 136 Thrailkill had pigeons play the role of matcher and mismatcher in the same session by splitting 137 each session into halves and reversing matcher-mismatcher roles between each half of the 7 138 session; Martin et al. did not do this. By creating a multiple schedule of games, Sanabria and 139 Thrailkill may have produced interaction between components. Such an outcome is often seen in 140 pigeon experiments where multiple schedules are used (e.g., Buck, Rothstein, & Williams, 141 1975). Moreover, we note that Martin et al. were scrupulous in ensuring procedural identity, 142 insofar as was practical, between the games played by chimpanzees and humans. Given our 143 speculation about multiple-schedule interaction and given that there are other procedural 144 differences between how Martin et al. and Sanabria and Thrailkill arranged their games, we 145 repeat pigeon game play in this report, this time mimicking Martin et al.’s design closely. 146 An expected result: Matching 147 Given the results of Sanabria and Thrailkill (2009), we are unsure pigeons will achieve 148 the NE in our experiment. However, there is another result we do expect: a high correlation 149 between changes in reinforcement ratios induced by various games and the choice ratios they 150 produce when these ratios are plotted in logarithmic space (Baum, 1974). This outcome defines 151 the Matching Law. 152 This is not a bold prediction. Give any animal, save humans (Lowe & Horne, 1985), two 153 manipulanda, each associated with a schedule of reinforcement, matching nearly always obtains 154 (Davison & McCarthy, 2016). In fact, the rapidity and regularity of matching relations in choice 155 has led some investigators to argue that matching in choice is innate (e.g., Gallistel, King, 156 Gottlieb, Balci, Papachristos, Szalecki, & Carbone, 2007). If our expectation of matching is 157 realized, and if pigeons also achieve the NE, interesting questions emerge depending on how the 158 reader explains matching. 159 160 One popular explanation is that matching is due to maximizing some dimension of reinforcement such as its rate or immediacy (Rachlin, Green, Kagel & Battalio, 1976; Shimp, 8 161 1966). To us, such a notion seems broadly compatible with the definition of the NE because the 162 NE defines a choice ratio where prospects for reinforcement cannot be improved upon unless 163 there is a change in a competitor’s choice allocation. In this case, achieving the NE and 164 matching likely share a common maximization process. Alternatively, some accounts of 165 matching attribute a matching outcome literally to a matching process (Herrnstein, 1970; 166 Gallistel et al., 2007). If this is the case, achieving the NE becomes an epiphenomenal correlate 167 of adherence to a matching process. 168 There is a long, contentious and unresolved literature on whether matching as outcome is 169 due to maximizing or a matching process. Regardless of which account is superior, explaining 170 achieving the NE in terms of matching seems advantaged because only this account could 171 accommodate choice in simple concurrent schedules as well as in game competition because 172 choice in concurrent schedules subsumes choice in competitive play. The converse is not the 173 case. This perspective raises the possibility that matching pennies in nonhuman animals may be 174 a competition in name only. Indeed, what they may perceive from a cognitive perspective may 175 not be a competition, but a standard concurrent schedule such as those seen in other matching 176 studies. 177 Experiment 1: Matching pennies in pigeons 178 Method 179 180 Subjects Four, one-year-old male Silver King pigeons that were used in a prior version of this 181 experiment served as subjects. This earlier experiment presented single-hopper presentations 182 that differed in their durations. Unfortunately, the design of the grain hopper failed to ensure 183 long hopper durations provided continuous access to grain. For this reason, multiple hopper 9 184 presentations of briefer duration were in the present version of the experiment. This permitted 185 grain to flow to the hopper opening at a constant rate. 186 Pigeons were maintained at 80% of their free-feeding weights and received supplemental 187 feeding of mixed grains when their weights fell below this level. They lived in an animal colony 188 where a 14-h light, 10-h dark cycle was maintained (lights on at 8 AM). Water was freely 189 available in their home cages. Sessions were conducted five days per week, usually in the 190 morning. 191 Apparatus 192 Training and testing occurred in two Med Associates (St. Albans, VT) operant pigeon 193 chambers (55.8 x 55.8 x 40.6 cm) that were enclosed in sound-attenuation chests. The chambers 194 had front and rear aluminum walls, Plexiglas side walls and ceiling, and a floor composed of 195 stainless-steel bars. Three evenly spaced cue lights were mounted in the front wall 10 cm above 196 the floor. Below the center cue light was a 2- by 3-cm (height by length) oval opening to a 197 mixed-grain food dispenser, the lower lip of which was 2.5 cm above the floor. A shielded, 100- 198 mA house light was mounted on the ceiling at the front of the chamber. Three 2.5-cm keys that 199 could be transilluminated from behind the front wall were arrayed 5.5 cm apart, center to center, 200 21.5 cm above the chamber floor. 201 Procedure 202 Pigeon pairs, one rewarded as a matcher and the other as a mismatcher, were exposed in 203 paired chambers to the symmetric, asymmetric and inspection games with reinforcer-amount 204 ratios conforming with the values described earlier for each of these games. The individual 205 reinforcer amount was 3 s of access to mixed grain. Different reinforcer-amount ratios were 206 created by multiple presentations of the hopper with 2 s intervening between successive 10 207 presentations. In the symmetric game, matches resulted in a single hopper presentation; in the 208 asymmetric game that followed, left-side matches produced three hopper cycles and right-side 209 matches produced one; and in the inspection game, left-side matches produced four hopper 210 cycles vs. one for right-side matches. For mismatchers in these games, correct mismatches 211 yielded one hopper cycle in the symmetric game and two in the other two games regardless of 212 the side chosen. 213 At the start of each session, the house light and center key light were illuminated with 214 white light. When one pigeon pecked the illuminated center key, it extinguished. When the other 215 pigeon pecked its center key, its light extinguished and then the side keys in both chambers were 216 illuminated with white light. If both pigeons pecked the same side key, the matcher earned a 217 reward. If the pigeons pecked opposite side keys, the mismatcher earned a reward. Once both 218 pigeons pecked a side key, the grain hopper provided the reward associated with those choices, 219 and the house light for the rewarded pigeon was extinguished for the duration of the hopper 220 cycle. A 30-s intertrial interval followed each presentation of a reward cycle. The center key 221 light was then illuminated again for the next trial unless the 100-trial, session-ending 222 contingency had been met. 223 Pigeons were randomly assigned to the roles of matcher and mismatcher. They then 224 played the symmetric game, followed by the asymmetric game, and finally the inspection game. 225 The pigeons were then switched to the opposite role and played the three games again in the 226 same order. Each game lasted no fewer than 12 sessions and no more than 25 sessions. A game 227 ended between those numbers if both pigeons in a pair had no data varying more than 15% from 228 the moving average of the last five sessions. All data are based on the last 200 trials of a game. 229 Results and Discussion 11 230 231 Pigeons approximate NE in matching pennies Figure 1 portrays the outcomes during the last 200 trials for the humans and chimpanzees 232 in Martin et al. (2014) and for the pigeons in this experiment. The data from Martin et al. were 233 digitized from Figures S5 and S6 from their supplemental online data set by using Plot Digitizer 234 version 2. The Figure presents the probability of a right choice by the mismatcher as a function 235 of the probability of a right by the matcher. The symmetric, asymmetric and inspection games 236 are arrayed in three panels from top to bottom. The NE, and preference data for the average 237 pigeon, chimpanzee and human are identified by different symbols (see key in bottom panel). 238 The error bars through the pigeon data define one standard error of the mean. To a first 239 approximation, it looks like humans in the inspection game (bottom panel) are farther from the 240 NE than pigeons and chimpanzees, and that averaged across all games, pigeons do as well as 241 chimpanzees. 242 Figure 2 tests this conclusion by quantifying the Euclidean distance from the NE across 243 all games for all species. As is apparent in the inspection game, the only game in which all 244 species are compared, humans do worse than pigeons and chimpanzees; and to our eyes, the 245 average chimpanzee’s performance is like that of the average pigeon (right pair of bars in Figure 246 2). However, it should be noted that averaging masks a possible between-game species 247 difference: chimpanzees more closely approximated the NE in the asymmetric game whereas 248 pigeons performed better than chimpanzees in the inspection game. These results are not 249 amenable to statistical test because Martin et al. (2014) only present population averages. 250 Martin et al. (2014) used a runs test to measure how closely chimpanzees and humans 251 adhered to the optimal pattern of stochastic response emission. The purpose of this measure was 252 to address the possibility that a player achieves the NE in terms of preference level even though 12 253 the player performed poorly because of how response sequences occurred. For example, in the 254 symmetric game, optimal performance is to respond stochastically with half of the responses 255 being to each alternative. But what if one player responded, say, ten times to the left alternative 256 and then ten times to the right alternative to achieve the NE preference-level statistic? In such a 257 case, preference level would not properly characterize the inadequacy of this response pattern 258 because surely such response sequencing would be co-opted by a competitor. 259 We did not collect runs data in this study, and therefore cannot use it to test for stochastic 260 choice allocation in game play. In our view, such a test lacks discriminative power (see Hachiga 261 & Sakagami, 2010). To illustrate the problem, imagine a player in the symmetric game achieved 262 the NE over 20 choices by responding twice to an alternative before switching. Such a pattern 263 would generate ten runs over 20 choices. It would pass the statistical expectation for stochastic 264 choice and could also achieve the NE even though choice was obviously patterned. Instead, we 265 have created an alternative statistic—payoffs per trial—to evaluate whether successive choices 266 were approximately independent. The Appendix shows how this statistic was calculated for all 267 three games based on the notion that choice is perfectly stochastic. For all games, Table 1 268 compares for each pigeon as a matcher and a mismatcher the expected number of payoffs per 269 trial vs. the number obtained. As is apparent, the differences between rewards received and 270 rewards expected by probabilistic responding were minor. These results support the view that 271 pigeons approximated the NE with successive choices being largely independent of each other. 272 This comparison is presented graphically in Figure 3. This figure shows that most of the 273 differences are small between the number of payoffs predicted by stochastic responding vs. those 274 obtained. Based on a paired t test (df = 47), there is no difference between the mean expected 275 and obtained payoffs per trial (-.0004). The 95% confidence interval is -.022 to .021. Despite 13 276 this outcome, we acknowledge that the force of our argument is weakened by our failing to 277 record runs data as done by Martin et al. (2014). Had those data been collected, it would have 278 been possible to compare directly sequential dependencies as measured by runs between our 279 study and theirs and assess whether any differences appeared in this measure between Martin et 280 al. and the present report. 281 One way to defend Martin et al.’s (2014) parallel cognitive-advantage and cognitive- 282 tradeoff hypotheses would be to deny our claim pigeons were in a competitive game analogous 283 to that given to Martin et al.’s chimpanzees. In Martin et al., chimpanzees were in the same 284 enclosure and each could turn its head to watch how the other chimpanzee responded and, in 285 principle, could use that information to make their matching or mismatching choices. Such an 286 option was unavailable to the competing pigeons because each was in an opaque enclosure. 287 Chimpanzees could see they were competing; pigeons could not. While this difference could 288 have mattered, it manifestly did not: Neither matching nor mismatching chimpanzee used the 289 choice of its competitor in a trial to guide its choice; had either done so, it would have created a 290 different data set, one where the informed subject made few errors and the other made many. 291 A final between-study difference warrants mention. Martin et al. (2014) cued the choice 292 of each chimpanzee’s opponent after each trial by flashing the opponent’s choice on each 293 touchscreen. We viewed such cuing as unnecessary because trial outcome (food or no food) 294 provided the same information as the flashing light in Martin et al. Many studies show that when 295 a discriminative stimulus is redundant, it does not control behavior (e.g., Reynolds, 1961). For 296 this reason, choices of each pigeon’s competitor were not signaled as done in Martin et al. 297 Pigeons approximate predictions of Matching Law 14 298 We have shown that pigeons, like chimpanzees, approximately achieve the NE in 299 matching pennies. In addition, we predicted pigeon performances, and by implication, 300 chimpanzee performances might also be described as due to equating choice ratios to the ratios 301 of the food amounts those choices produce (matching). To test this prediction, we need target 302 studies from the matching literature where food amounts and payoff likelihoods have been varied 303 between two alternatives. We use Schneider (1973) for this purpose. In Schneider’s study, 304 pigeons chose between two keys, each of which was associated with a Variable-Interval (VI) 305 schedule that, depending on the condition, delivered different ratios of food pellets occasionally 306 for a choice. He found that preference depended on both reinforcer rate and reinforcer 307 magnitude (number of pellets) delivered for each alternative. He represented choice behavior in 308 terms of the following generalized matching equation: 309 log (R1/R2) = a log (r1/r2) + b log (q1/q2) + c 310 where R, a, r, b, q and c respectively stand for responses (R), a parameter for reinforcer- 311 frequency ratios (a), reinforcement frequency (r), a parameter for reinforcer amount ratios (b), 312 quantity of food per reinforcer (q), and bias (c). The subscripts 1 and 2 distinguish between the 313 choice alternatives. The a and b parameters accommodate slope differences in the plot whereas 314 the c parameter accommodates differences in Y-intercept values. (1) 315 The line of best fit that results from applying Equation 1 to individual pigeon 316 performances in Experiment 1 is presented in Figure 4. Do the matching-pennies data from 317 Martin et al. (2014) and Sanabria and Thrailkill (2009) look like the data in Figure 4, showing 318 matching-like outcomes? 319 320 Figure 4 is based on the actual choice-ratio, reinforcer-frequency and reinforcer-amount data for Experiment 1. To evaluate Martin et al. (2014) and Sanabria and Thrailkill (2009) in 15 321 terms of the same measures requires the simplifying assumption that all choice is stochastic in 322 these two studies. The aggregate amount of reinforcement was estimated from choice 323 probabilities. To calculate the expected reinforcement frequency to the left and right keys for the 324 matcher, the respective frequency of left and right choices by the matcher was multiplied by the 325 respective frequency of left and right choices by the mismatcher. These data were used to 326 calculate a left-key/right-key reinforcer ratio. A similar exercise was done in the case of the 327 mismatcher, accommodated to reflect that reinforcers for the mismatcher require choice 328 mismatches. 329 To ensure comparability to our pigeon results, we make the same assumptions for that 330 data set. Figure 5 presents data based on these estimates for Experiment 1 (top panel), Martin et 331 al. (middle panel) and Thrailkill and Sanabria (bottom panel). All three data sets look like data 332 from choice matching studies. 333 Experiment 2: Application of the Cumulative-Effects Model to Matching Pennies 334 We have argued that chimpanzees and pigeons may view matching pennies not as a 335 competitive game, but as a choice procedure akin to those that populate the operant literature on 336 concurrent performances. Certainly, the fact that approximating the NE in a matching-pennies 337 games also approximates matching relations raises this as a possibility. Arguing that 338 chimpanzees in Martin et al. (2014) were simply adhering to the Matching Law does damage to 339 their claims of a chimpanzee cognitive advantage in matching pennies because many species 340 throughout the animal kingdom match. Presumably, many are less cognitively gifted than 341 chimpanzees. And based on what we have seen so far, it may well be the case that animals that 342 match also achieve the NE. 16 343 In this second experiment, we demonstrate how a cognitively simple matching model 344 from the operant choice literature succeeds in achieving the NE in matching-pennies games. 345 This result complements the analytic data from Experiment 1 by now synthesizing the NE 346 outcome of chimpanzees by means of the Matching Law. 347 There are several models to choose from that predict matching. For example, the copyist 348 model (Tanno & Silberberg, 2012) simulates matching even though it posits that an animal 349 simply reproduces what was reinforced before. In principle, it could be used to test whether it 350 also predicts the NE in matching-pennies games. Nevertheless, we have selected the cumulative- 351 effects model for simulation (Davis, Staddon, Machado, & Palmer, 1993) because it produces 352 matching in choice, and it is easier to program on a computer playing matching-pennies games. 353 The cumulative-effects model posits that an animal calculates the number of reinforcers 354 provided by a choice alternative divided by the number of responses made to produce them. The 355 alternative which has the higher reinforcers-per-response value garners the next response. Davis 356 et al. (1993) demonstrated that this decision rule produces matching. To test whether it also 357 achieves the NE in the Martin et al. (2014) games, twelve 400-trial matching-pennies games 358 were played by matcher/mismatcher stat animals that followed the cumulative-effects model in 359 the symmetric, asymmetric and inspection games. 360 The stat animals were programmed to begin a simulation by responding with equal 361 probability to each alternative. Once both alternatives had delivered simulated reinforcers to 362 both stat animals according to the requirements of a given game, a reinforcers-per-response 363 statistic was calculated for each alternative, and each subsequent choice went to the simulated 364 key with the higher value. To illustrate the operation of this simulation, imagine that up to the 365 current trial the matcher had obtained 160 reinforcements from making 200 responses on the left 17 366 key (reinforcement rate = 160/200 or 0.8), the key that provides a reinforcer four times larger 367 than that provided by the right key. If the next left response produced a reward, the left-key 368 reinforcement rate was updated to 164/201 or 0.82. On the other hand, if the matcher chose the 369 right key and got a four-fold smaller reward, the numerator of the right-key reinforcement rate 370 was only increased by one. 371 Figure 6 presents the outcomes of these simulations in the format used in Figure 1. As is 372 apparent, the cumulative-effects model does an excellent job of achieving the NE in the games 373 used by Martin et al. (2014). These outcomes support the view that chimpanzees and pigeons 374 may achieve the NE by following a simple matching rule. 375 Figure 7 produces the matching data (top panel) and expected-payoff data (bottom panel) 376 using the axes of Figures 4 and 3, respectively. As is apparent, simulated performances are 377 consistent with matching predictions and the expected-payoff data, generated by perfectly 378 stochastic choice allocation, look quite like those seen in real pigeons. The mean difference 379 between the expected and obtained payoffs/trial was .003. The 95% confidence interval was 380 from -.008 to .014. 381 382 General Discussion Experiment 1 shows that pigeons and chimpanzees are approximately equivalent in their 383 capacity to achieve the NE in the competitive games used by Martin et al. (2014), and that both 384 species do better in this task than humans. Martin et al.’s theses explaining the superiority of 385 chimpanzee-to-human game play are now in question unless the reader extends these notions to 386 explain the superiority of pigeon-to-human game play. Moreover, choice allocation in our 387 pigeon study was well described by a matching equation. In Experiment 2 we showed by 388 simulation that a model that produces matching in concurrent schedules also achieves the NE 18 389 when applied to matching-pennies games. Such an outcome may mean that matching-pennies 390 games in nonhumans should not be viewed as competitions to be evaluated by proximity to the 391 NE. Instead, they are choice procedures that should be described in terms of the Matching Law. 392 These outcomes invite consideration of whether achieving the NE should be viewed as 393 due to matching as process or that both matching and achieving the NE are derived from some 394 other behavioral principle such as reinforcement maximization. As noted earlier, stochastically 395 allocated choice produces matching in standard concurrent schedules when the choice ratio 396 approximates matching relations (Rachlin et. al., 1976). This outcome seems compatible with 397 the definition of the NE as the choice ratio that optimizes income unless competitor preferences 398 change. A second possibility is that matching itself is responsible for achieving the NE, and the 399 kinds of measures used by NE theorists fail to evidence this fact. This argument gains credence 400 if the reader accepts the arguments made by Gallistel et al. (2007) that matching as process is 401 innate. In either case, the third possibility to consider—that pursuit of the NE accommodates 402 concurrent-schedule performances and competitive game play—does not seem credible. The 403 obvious problem is that on conventional concurrent schedules, subject choice does not face a 404 competitor. 405 Summary, implications and conclusions 406 In this report, we provide data relevant to Martin et al.’s (2014) hypotheses regarding 407 superiority in matching pennies by chimpanzees by showing: (a) chimpanzees are likely hardly 408 unique in achieving the NE in matching-pennies games; (b) what is viewed as a competitive 409 game by Martin et al. (2014) may, in terms of its psychology, be closer to conventional 410 concurrent schedules as used in behavior analysis; and (c) chimpanzees and pigeons achieving 411 the NE in matching pennies may be an epiphenomenal correlate of the operation of a matching 19 412 rule such as that provided by the cumulative-effects model (alternatively both outcomes may be 413 due to another process such as reinforcement maximization). In any case, the finding of 414 matching in pigeons in matching pennies (Experiment 1) that can be synthesized by simulation 415 (Experiment 2) supports the expectation that many species should achieve the NE in matching 416 pennies because virtually all species tested to date match choice ratios to reinforcer ratios. 417 If the reader accepts our view, the findings in this report weaken confidence in the notion 418 of Martin et al. (2014) that chimpanzees have task-specific cognitive advantages over humans 419 and possibly many other species in playing matching-pennies games. In our view, Martin et al.’s 420 account is one-sided for advancing unusual explanations for superior game play in chimpanzees 421 in the face of an obvious alternative explanation for their data: Humans perform more poorly not 422 because chimpanzees are cognitively advantaged in matching pennies or because humans have 423 traded off cognitive capacity for language, but because of the inherent greater complexity of 424 human mental function. 425 Evidence of human mental complexity compromising human performances abounds in 426 prospect theory where we see many examples of cognitive errors unlikely to be found in other 427 animals (Kahneman & Tversky, 1979). Prospect theorists do not advance these errors as 428 examples of how humans are more limited than other animals. Although they are typically silent 429 about nonhumans, we think it likely they would attribute the results of Martin et al. (2014) not to 430 the limits of human mental function when compared to other animals, but to their expanse. 431 The literature on probability matching is a classic illustration of where humans do more 432 poorly than other animals. In a probability-matching task, a reward is assigned with unequal 433 probability to each of two choices. Humans often behave sub-optimally by matching their 434 relative choices to the relative outcomes they produce. For example, if a random 70% of the 20 435 rewards are given for the choice “left” and the rest are given for the choice “right”, humans will 436 tend to choose “left” 70% of the time. Few other species tested on this task make this error— 437 virtually all their choices are “left” (Fantino & Esfaniari 2002; Graf, Bullock, & Bitterman, 438 1964; Wilson, 1960). To our knowledge, no probability-matching theorist advances the idea that 439 humans do more poorly than other animals because of their cognitive limitations. Instead, they 440 point to added features of human mental function such as the gambler’s fallacy (Gold, 1997) 441 which cause human performance to be inferior (e.g., Tversky & Kahneman, 1974). Data from 442 Derks and Paclisanu (1967) support this interpretation. They showed through age five, children 443 maximize on probability-matching tasks, just like nonhuman animals; only when older do they 444 probability match. If one accepts as plausible that human adults are more cognitively advanced 445 than children, it seems reasonable to interpret maximizing to simpler mental processes evident in 446 children and probability matching to a cognitive error such as the gambler’s fallacy that is 447 present in adults. 448 If we attribute inferior matching-pennies performance to the greater complexity of human 449 mental function, predictive parsimony obtains. For example, no longer is explanation burdened 450 by the fact that accepting chimpanzees have a parallel cognitive advantage leads to the 451 implication that pigeons also have a parallel cognitive advantage. Instead, this perplexing 452 outcome for theory disappears: Human mental sophistication causes humans to perform more 453 poorly in matching pennies than pigeons. It is also resolves the singular status of Martin et al.’s 454 (2014) data being outside the Dennett (1983) dichotomy. They are not. 455 456 457 21 458 459 460 461 References Baum, W. M. (1974). On two types of deviation from the matching law: Bias and undermatching. Journal of the Experimental Analysis of Behavior, 22, 231-242. Buck, S. L., Rothstein. B., & Williams, B. A. (1975). A re-examination of local contrast in 462 multiple schedules. Journal of the Experimental Analysis of Behavior, 24, 291-301. 463 Davis, D. G., Staddon, J. E., Machado, A., & Palmer, R. G. (1993). The process of recurrent 464 465 466 467 468 469 470 471 472 473 choice. Psychological Review, 100, 320-341. Davison, M., & McCarthy, D. (2016). The matching law: A research review. London, England: Routledge. Dennett, D. C. (1983). Intentional systems in cognitive ethology: The “Panglossian paradigm” defended. Behavioral and Brain Sciences, 6, 343-355. Derks, P. L., & Paclisanu, M. I. (1967). Simple strategies in binary prediction by children and adults. Journal of Experimental Psychology, 73, 278-285. Fantino, E., & Esfandiari, A. (2002). Probability matching: Encouraging optimal responding in humans. Canadian Journal of Experimental Psychology, 56, 58-63. Gallistel, C. R., King, A. P., Gottlieb, D., Balci, F., Papachristos, E. B., Szalecki, M., & Carbone, 474 K. S. (2007). Is matching innate? Journal of the Experimental Analysis of Behavior, 87, 475 161-199. 476 477 478 479 Genty, E., Karpel, H., & Silberberg, A. (2012). Time preferences in long-tailed macaques (Macaca fascicularis) and humans (Homo sapiens). Animal Cognition, 15, 1161-1172. Gold, E. (1997). The gambler’s fallacy (Doctoral Dissertation). Carnegie Mellon University, Social and Decision Sciences Department. 22 480 481 482 483 484 485 486 487 488 489 490 491 492 Graf ,V., Bullock, D. H., & Bitterman, M. E. (1964). Further experiments on probabilitymatching in the pigeon. Journal of the Experimental Analysis of Behavior, 7, 151-157. Hachiga, Y., & Sakagami, T. (2010). A runs-test algorithm: Contingent reinforcement and response run structures. Journal of the experimental analysis of behavior, 93, 61-80. Herrnstein, R. J. (1970). On the law of effect. Journal of the Experimental Analysis of Behavior, 13, 243-266. Inoue, S., & Matsuzawa, T. (2007). Working memory of numerals in chimpanzees. Current Biology, 17, R1004-R1005. Jensen, K., Call, J., & Tomasello, M. (2007). Chimpanzees are rational maximizers in an ultimatum game. Science, 318, 107-110. Kahneman, D., & Tversky, A. (1979). Prospect theory: An analysis of decision under risk. Econometrica, 47, 263-291. Lowe, C. F., & Horne, P. J. (1985). On the generality of behavioural principles: Human choice 493 and the matching law. In C. F. Lowe, M. Richelle, D. E. Blackman & C. M. Bradshaw 494 (Eds.). Behaviour analysis & contemporary psychology (pp. 97-115). London, England: 495 Lawrence Erlbaum Associates. 496 Martin, C. F., Bhui, R., Bossaerts, P., Matsuzawa, T., & Camerer, C. (2014). Chimpanzee choice 497 rates in competitive games match equilibrium game theory predictions. Scientific 498 Reports, 4, 5182. 499 Rachlin, H., Green, L., Kagel, J. H., & Battalio, R. C. (1976). Economic demand theory and 500 psychological studies of choice. In Psychology of Learning and Motivation (Vol. 10, pp. 501 29-154). Cambridge, MA: Academic Press. 23 502 503 504 Reynolds, G. S. (1961). Attention in the pigeon. Journal of the Experimental Analysis of Behavior, 4, 203-208. Rosati, A. G., Stevens, J. R., Hare, B., & Hauser, M. D. (2007). The evolutionary origins of 505 human patience: temporal preferences in chimpanzees, bonobo and human adults. 506 Current Biology, 17, 1663-1668. 507 Sanabria, F., & Thrailkill, E. (2009). Pigeons (Columba livia) approach Nash equilibrium in 508 experimental matching pennies competitions. Journal of the Experimental Analysis of 509 Behavior, 91, 169-183. 510 Schneider, J. W. (1973). Reinforcer effectiveness as a function of reinforcer rate and magnitude: 511 A comparison of concurrent performances. Journal of the Experimental Analysis of 512 Behavior, 20, 461-471. 513 514 515 516 517 518 519 520 521 522 523 524 Shimp, C. P. (1966). Probabilistically reinforced choice behavior in pigeons. Journal of the experimental analysis of behavior, 9, 443-455. Silberberg, A, & Kearns, D. (2009). Memory for the order of briefly presented numerals in humans as a function of practice. Animal Cognition, 12, 405-407. Smith, P., & Silberberg, A. (2010). Rational maximizing by humans (Homo sapiens) in an ultimatum game. Animal Cognition, 13, 671-677. Tanno, T., & Silberberg, A. (2012). The copyist model of response emission. Psychonomic Bulletin & Review, 19, 759-778. Tversky, A., & Kahneman, D. (1974). Judgment under uncertainty: Heuristics and biases. Science, 185, 1124–1131. Wilson, W. A. (1960). Two-choice behavior of monkeys. Journal of Experimental Psychology, 59, 207-208. 24 525 Appendix 526 Suppose p is the left-choice probability of the mismatcher and q is the left-choice probability of 527 the matcher. The expected number of reinforcers per trial in each game was calculated as 528 follows: 529 Symmetric Game 530 For the matcher, p * 1 + (1 - p) * 0 = p for Left, and p * 0 + (1 – p) * 1 = 1 – p for Right. 531 For the mismatcher, q * 0 + (1 – q) * 1 = 1 – q for Left, and q * 1 + (1 – q) * 0 = q for Right. 532 Asymmetric Game 533 For the matcher, p * 3 + (1 – p) * 0 = 3p for Left and p * 0 + (1 – p) * 1 = 1 – p for Right. 534 For the mismatcher, q * 0 + (1 – q) * 2 = 2 – 2q for Left and q * 2 + (1 – q) * 0 = 2q for Right. 535 Inspection Game 536 For the matcher, p * 4 + (1 – p) * 0 = 4p for Left and p * 0 + (1 – p) * 1 = 1 – p for Right. 537 For the mismatcher, q * 0 + (1 – q) * 2 = 2 – 2q for Left and q * 2 + (1 – q) * 0 = 2q for Right. 538 539 540 541 542 543 544 545 546 547 548 549 25 550 551 552 Table 1 26 553 554 Figure Captions Figure 1. Probability of a right in mismatchers as a function of the probability of a right 555 in matchers in three different competitive games. Pigeon data are from this experiment. All 556 other data are from Martin et al. (2014). See text for other details. 557 Figure 2. Distance from the NE for pigeons, chimpanzees and humans in symmetric 558 game, asymmetric game and inspection game. Chimpanzee and human data are from Martin et 559 al. (2014). Bars on far, right side of panel are all-game averages for pigeons and chimpanzees. 560 Figure 3. A graphic presentation of the results in Table 1. This figure presents the 561 relative frequency of the differences between the expected payoff per trial that would be 562 produced by stochastic response emission vs. the results obtained based on pigeon choice 563 frequencies. 564 Figure 4. The panel’s X and Y axes respectively present log choice ratios as a function of 565 the log ratio of the total amount of food each choice alternative provided in a session. Individual 566 performances of pigeons in each role (matcher or mismatcher) in each type of matching-pennies 567 game from Experiment 1 populate the graph. The best-fit function is based on Equation 1. The 568 R2 statistic in the panel defines the proportion of the individual-subject data variance 569 accommodated by the best-fit function. 570 Figure 5. Matching analysis of the results of the current experiment and those for 571 chimpanzees from Martin et al. and for pigeons from Sanabria and Thrailkill based on Equation 572 1. See text for other details. 573 Figure 6. The probability of a right in mismatchers as a function of the probability of a 574 right in matchers in three different matching-pennies games based on the cumulative-effects 575 model. In all games, the closed circles representing simulated animal performances overlap 27 576 with the NE (open squares). Error bars through these data points define one standard error of the 577 mean. Mean performances for chimps from Martin et al. (2014) are also presented. 578 Figure 7. Log ratio of choices to each key in the cumulative-effects model simulations 579 for all games from Martin et al. for matchers and mismatchers (top panel). The bottom panel 580 presents the relative frequency distribution of the differences between expected and obtained 581 payoffs per trial in these simulations. 582 583 584 585 586 587 588 589 590 591 592 593 594 595 596 597 598 28 599 600 601 Figure 1 29 602 603 604 Figure 2 30 605 606 607 608 Figure 3 31 609 610 611 612 613 Figure 4 32 614 615 Figure 5 33 616 617 618 619 Figure 6 34 620 621 Figure 7