1 00:00:00,120 --> 00:00:01,800 The following content is provided 2 00:00:01,800 --> 00:00:04,040 under a Creative Commons license. 3 00:00:04,040 --> 00:00:06,880 Your support will help MIT OpenCourseWare continue 4 00:00:06,880 --> 00:00:10,740 to offer high quality educational resources for free. 5 00:00:10,740 --> 00:00:13,500 To make a donation or view additional materials from 6 00:00:13,500 --> 00:00:17,270 hundreds of MIT courses, visit MIT OpenCourseWare 7 00:00:17,270 --> 00:00:21,318 at ocw.mit.edu . 8 00:00:21,318 --> 00:00:22,550 PROFESSOR: OK. 9 00:00:22,550 --> 00:00:28,840 So we've started talking about many of the key concepts 10 00:00:28,840 --> 00:00:32,930 in behavioral studies of learning. 11 00:00:32,930 --> 00:00:36,750 And I'm basing this on Konrad Lorenz. 12 00:00:36,750 --> 00:00:47,360 And this is where we ended. 13 00:00:47,360 --> 00:00:50,100 We were about to talk about avoidance responses acquired 14 00:00:50,100 --> 00:00:52,120 through trauma. 15 00:00:52,120 --> 00:00:56,200 We had just talked about conditioned reflexes. 16 00:00:56,200 --> 00:01:00,180 Remember, he sees the conditioned reflex, 17 00:01:00,180 --> 00:01:04,860 as the way it has been studied in experimental situations, 18 00:01:04,860 --> 00:01:06,580 as a kind of stimulus selection. 19 00:01:09,100 --> 00:01:10,990 And that's always the way it's interpreted. 20 00:01:10,990 --> 00:01:13,410 And there are many, many experiments 21 00:01:13,410 --> 00:01:19,480 that perfectly fit the Lorenz definition, even 22 00:01:19,480 --> 00:01:22,210 though Pavlov's original experiment 23 00:01:22,210 --> 00:01:28,650 that so popularly described as an example 24 00:01:28,650 --> 00:01:31,360 of classical conditioning. 25 00:01:31,360 --> 00:01:33,860 So we'll bring that up again a little later because you'll 26 00:01:33,860 --> 00:01:36,780 see, then, what Lorenz calls it. 27 00:01:36,780 --> 00:01:39,200 But first let's finish talking about learning 28 00:01:39,200 --> 00:01:43,480 through association without feedback, without feedback 29 00:01:43,480 --> 00:01:48,230 reporting the success of the movement. 30 00:01:48,230 --> 00:01:52,380 So we'll talk about avoidance response 31 00:01:52,380 --> 00:01:53,750 acquired through trauma. 32 00:01:53,750 --> 00:01:55,950 It's very interesting kind of learning, 33 00:01:55,950 --> 00:01:59,990 because it's pretty irreversible. 34 00:01:59,990 --> 00:02:03,750 It's another type of stimulus selection. 35 00:02:03,750 --> 00:02:06,590 It can occur after just one experience, 36 00:02:06,590 --> 00:02:10,740 and then be considerably permanent. 37 00:02:10,740 --> 00:02:15,750 He talks about how trainers of dogs and horses-- 38 00:02:15,750 --> 00:02:24,140 just like psychoanalysts-- they know how, once an animal learns 39 00:02:24,140 --> 00:02:27,720 to avoid something, some stimulus in his environment 40 00:02:27,720 --> 00:02:33,750 becomes associated with avoidance, something that's 41 00:02:33,750 --> 00:02:37,540 frightened the animal a lot, or frightened the person, 42 00:02:37,540 --> 00:02:40,170 becomes associated with certain stimuli, 43 00:02:40,170 --> 00:02:42,345 it's very difficult to change that. 44 00:02:45,550 --> 00:02:47,730 In the laboratory usually it's done 45 00:02:47,730 --> 00:02:51,830 by pairing sound with electric shock. 46 00:02:51,830 --> 00:02:56,430 Sometimes it's not sound, it's another stimulus. 47 00:02:56,430 --> 00:03:00,280 And then later, with later presentations of the sound, 48 00:03:00,280 --> 00:03:04,785 you will get autonomic responses, increased heart rate 49 00:03:04,785 --> 00:03:06,500 and blood pressure. 50 00:03:06,500 --> 00:03:10,280 And at least some forms of it are just 51 00:03:10,280 --> 00:03:12,950 one trial learning, one experience. 52 00:03:12,950 --> 00:03:15,870 The most common type in the laboratory, as many of you 53 00:03:15,870 --> 00:03:18,250 know, is what we call the step down response, 54 00:03:18,250 --> 00:03:21,560 because a little rodent on a raised platform 55 00:03:21,560 --> 00:03:26,920 will tend to step down from a platform, unless his vision 56 00:03:26,920 --> 00:03:30,190 indicates that he's very high up, 57 00:03:30,190 --> 00:03:33,220 then he's much less likely to step down. 58 00:03:33,220 --> 00:03:37,210 But if he can step down fairly comfortably he will do it. 59 00:03:37,210 --> 00:03:40,500 But when he steps down, the grid-- 60 00:03:40,500 --> 00:03:44,060 he steps on an electrified grid-- then, 61 00:03:44,060 --> 00:03:46,670 just in that one trial, you bring him back later, 62 00:03:46,670 --> 00:03:51,700 and he'll just stay and stay and stay on the platform. 63 00:03:51,700 --> 00:03:55,790 So that's an example of avoidance responses acquired 64 00:03:55,790 --> 00:03:56,415 through trauma. 65 00:03:58,940 --> 00:04:05,920 And I just point out here that most psychologists-- anything 66 00:04:05,920 --> 00:04:07,730 I've added in red on these slides 67 00:04:07,730 --> 00:04:10,990 is not in the thing posted online, 68 00:04:10,990 --> 00:04:15,030 but I'll change that to the annotated version later on. 69 00:04:15,030 --> 00:04:17,620 So most psychologists include this kind 70 00:04:17,620 --> 00:04:19,940 of learning in the classical conditioning category. 71 00:04:19,940 --> 00:04:24,370 But it's got enough differences, as Lorenz points out, 72 00:04:24,370 --> 00:04:25,250 to separate it. 73 00:04:28,710 --> 00:04:32,380 In other words, animals appear to have evolved differences. 74 00:04:32,380 --> 00:04:35,770 That's why this type is so irreversible compared 75 00:04:35,770 --> 00:04:40,984 with other types of stimulus selection learning. 76 00:04:40,984 --> 00:04:43,275 And then imprinting, which we have talked about before. 77 00:04:47,310 --> 00:04:50,170 I said here-- this is on the posted slides, 78 00:04:50,170 --> 00:04:54,320 both of the previous two-- that is, both avoidance 79 00:04:54,320 --> 00:04:57,050 response acquired through trauma and conditioned reflexes 80 00:04:57,050 --> 00:04:59,320 are irreversible. 81 00:04:59,320 --> 00:05:02,090 It's certainly true of the avoidance responses acquired 82 00:05:02,090 --> 00:05:02,780 through trauma. 83 00:05:02,780 --> 00:05:04,560 Relatively speaking, it's irreversible. 84 00:05:07,490 --> 00:05:09,470 Other types of classical conditioning 85 00:05:09,470 --> 00:05:11,010 are perhaps less irreversible. 86 00:05:14,600 --> 00:05:17,690 But we know that imprinting will cause 87 00:05:17,690 --> 00:05:21,390 a fixation of specific responses. 88 00:05:21,390 --> 00:05:22,680 They're very specific. 89 00:05:22,680 --> 00:05:27,610 We've learned that birds don't become imprinted on humans. 90 00:05:27,610 --> 00:05:28,700 There's no such thing. 91 00:05:28,700 --> 00:05:32,400 They become imprinted for particular situations, 92 00:05:32,400 --> 00:05:38,610 particular kind of responses, like his jackdaws. 93 00:05:38,610 --> 00:05:40,940 Because of early imprinting, they 94 00:05:40,940 --> 00:05:47,060 can become fixated on companions for sex, 95 00:05:47,060 --> 00:05:50,590 or they can become separately imprinted on companions 96 00:05:50,590 --> 00:05:58,440 for flocking and flying, and separately for parental care. 97 00:05:58,440 --> 00:06:01,290 And these things are quite separable. 98 00:06:01,290 --> 00:06:04,260 And all that became clear in his studies of jackdaws. 99 00:06:04,260 --> 00:06:07,590 And it's also clear in geese. 100 00:06:10,660 --> 00:06:13,270 Geese that are raised with humans 101 00:06:13,270 --> 00:06:17,490 can become imprinted for what he calls 102 00:06:17,490 --> 00:06:19,760 filial and social responses. 103 00:06:19,760 --> 00:06:23,310 And yet in their sexual behavior they're not affected. 104 00:06:23,310 --> 00:06:26,720 They will have sexual behavior normally 105 00:06:26,720 --> 00:06:29,030 with other members of their species. 106 00:06:29,030 --> 00:06:32,097 And so there are real species differences in this. 107 00:06:36,530 --> 00:06:42,240 You can separate imprinting from conditioning with reward 108 00:06:42,240 --> 00:06:44,940 by simply pitting them against each other. 109 00:06:44,940 --> 00:06:48,910 For example, you can have animals 110 00:06:48,910 --> 00:06:57,090 that are imprinted on, say, humans as sexual partners. 111 00:06:57,090 --> 00:07:01,570 And we've had examples of that, especially in the jackdaws. 112 00:07:04,190 --> 00:07:06,390 But it's been done with a number of other species. 113 00:07:06,390 --> 00:07:10,390 And in his book, he describes specific species 114 00:07:10,390 --> 00:07:16,015 where a species was imprinted for mating on another species, 115 00:07:16,015 --> 00:07:19,060 a similar species but different. 116 00:07:19,060 --> 00:07:22,110 Well, if you deprive the animal for a long period of time, 117 00:07:22,110 --> 00:07:24,880 you never expose him anymore to that species 118 00:07:24,880 --> 00:07:29,520 he's imprinted on, he eventually will-- 119 00:07:29,520 --> 00:07:32,340 because his action-specific-- what happens? 120 00:07:32,340 --> 00:07:35,020 You deprive an animal for a very long time, 121 00:07:35,020 --> 00:07:36,640 the action-specific potential, that 122 00:07:36,640 --> 00:07:38,800 is, the internal drive becomes so high, 123 00:07:38,800 --> 00:07:42,710 thresholds are lowered, he's likely to start mating 124 00:07:42,710 --> 00:07:43,980 with its own species. 125 00:07:43,980 --> 00:07:44,480 OK. 126 00:07:44,480 --> 00:07:46,650 So you let him do that for a while, 127 00:07:46,650 --> 00:07:49,710 taking care of young, everything. 128 00:07:49,710 --> 00:07:54,710 And now you bring back the other species 129 00:07:54,710 --> 00:07:56,810 that he was imprinted on. 130 00:07:56,810 --> 00:07:58,390 What happens? 131 00:07:58,390 --> 00:08:03,670 Either immediately or gradually, he switches over, 132 00:08:03,670 --> 00:08:05,170 he switches back. 133 00:08:05,170 --> 00:08:07,630 With all the reward and everything associated 134 00:08:07,630 --> 00:08:11,540 with mating with his own species, 135 00:08:11,540 --> 00:08:17,710 he still will switch back to the imprint of species, which 136 00:08:17,710 --> 00:08:21,840 shows the independence of these two processes. 137 00:08:21,840 --> 00:08:25,090 Then he gives the example of innate releasing mechanisms 138 00:08:25,090 --> 00:08:29,150 in mallard males and females. 139 00:08:29,150 --> 00:08:31,255 The males can be sexually imprinted 140 00:08:31,255 --> 00:08:35,330 to other species, but not the females. 141 00:08:35,330 --> 00:08:37,860 The reason is they're so dominated 142 00:08:37,860 --> 00:08:42,630 by innate releasing mechanisms that 143 00:08:42,630 --> 00:08:46,740 respond to the mallard drake's colors, 144 00:08:46,740 --> 00:08:51,310 his mating colors, nuptial colors. 145 00:08:51,310 --> 00:08:56,240 So it's a very different situation for male and female. 146 00:08:56,240 --> 00:08:59,710 Just like there are differences from one species to the other, 147 00:08:59,710 --> 00:09:03,990 there can be differences between the two sexes. 148 00:09:03,990 --> 00:09:07,250 So and then finally what he calls conditioned inhibition. 149 00:09:09,760 --> 00:09:16,000 And we train domestic animals and circus animals. 150 00:09:16,000 --> 00:09:23,640 We even train tigers and bears for circus performance. 151 00:09:23,640 --> 00:09:30,870 And they learn, they are taught, to inhibit natural responses. 152 00:09:30,870 --> 00:09:33,710 Obviously you're going to be in a circus, 153 00:09:33,710 --> 00:09:37,990 you don't want them to be showing prey-catching responses 154 00:09:37,990 --> 00:09:41,290 and so forth to a little child running 155 00:09:41,290 --> 00:09:43,330 in front of the audience. 156 00:09:43,330 --> 00:09:49,400 So you can suppress, you can condition them 157 00:09:49,400 --> 00:09:52,990 so they're inhibiting their normal responses. 158 00:09:52,990 --> 00:09:56,490 The problem is, if the action-specific potentials 159 00:09:56,490 --> 00:10:02,380 build up and build up and build up because of long suppression, 160 00:10:02,380 --> 00:10:06,410 you can sometimes have very dangerous consequences. 161 00:10:06,410 --> 00:10:11,720 And his examples are pack animals, like wolves. 162 00:10:11,720 --> 00:10:17,690 A dominant pack member can totally suppress resistance 163 00:10:17,690 --> 00:10:19,520 by other members of the pack. 164 00:10:19,520 --> 00:10:21,810 And so you'll have passive members of the pack. 165 00:10:21,810 --> 00:10:26,130 But they're really-- if he's at the bottom of the totem 166 00:10:26,130 --> 00:10:28,980 pole he's really repressed. 167 00:10:28,980 --> 00:10:36,890 He can suddenly-- his desire to respond more 168 00:10:36,890 --> 00:10:41,590 normally and oppose that leader can suddenly break out, 169 00:10:41,590 --> 00:10:43,380 and basically all hell breaks loose 170 00:10:43,380 --> 00:10:46,729 and the animal becomes very dangerous, 171 00:10:46,729 --> 00:10:48,770 because the normal inhibitions can suddenly fail. 172 00:10:48,770 --> 00:10:52,910 And there are specific examples Lorenz goes through. 173 00:10:52,910 --> 00:10:56,710 Hugo Van Lawick studied this in African hunting dogs. 174 00:10:56,710 --> 00:10:58,770 And he showed examples of it. 175 00:10:58,770 --> 00:11:01,050 They're another group-living animal. 176 00:11:01,050 --> 00:11:05,320 There's a dominant animal, and the animals 177 00:11:05,320 --> 00:11:09,230 low down on the social hierarchy can occasionally 178 00:11:09,230 --> 00:11:13,010 do this if their normal behavior is suppressed for too long. 179 00:11:13,010 --> 00:11:14,900 We mentioned they're already in wolves. 180 00:11:14,900 --> 00:11:17,470 It happens in humans, too, in a situation 181 00:11:17,470 --> 00:11:23,840 that certainly has legal consequences, namely 182 00:11:23,840 --> 00:11:25,370 spousal murder. 183 00:11:25,370 --> 00:11:29,940 So it usually happens when a woman is so totally controlled 184 00:11:29,940 --> 00:11:35,580 by a spouse that her normal reactions are totally 185 00:11:35,580 --> 00:11:36,280 suppressed. 186 00:11:36,280 --> 00:11:40,680 Her normal drives she cannot express. 187 00:11:40,680 --> 00:11:44,150 And if this goes on and on and on, 188 00:11:44,150 --> 00:11:47,440 especially if she's cruelly treated, 189 00:11:47,440 --> 00:11:51,010 she eventually-- her inhibitions can suddenly fail, 190 00:11:51,010 --> 00:11:54,600 and she just kills him. 191 00:11:54,600 --> 00:11:59,670 And when that comes to the court, women in that situation 192 00:11:59,670 --> 00:12:03,240 are often acquitted. 193 00:12:03,240 --> 00:12:05,790 So this does have some recognition 194 00:12:05,790 --> 00:12:12,480 in our legal system, this consequences of basically 195 00:12:12,480 --> 00:12:16,110 conditioned inhibition, although in the legal sense 196 00:12:16,110 --> 00:12:17,070 it's not called that. 197 00:12:19,740 --> 00:12:24,430 So now, learning affected by the consequences of behavior. 198 00:12:24,430 --> 00:12:27,110 These are the common kinds. 199 00:12:27,110 --> 00:12:30,040 Conditioned appetitive behavior we'll deal with first. 200 00:12:30,040 --> 00:12:33,500 Conditioned aversion, conditioned action. 201 00:12:33,500 --> 00:12:35,310 Conditioned aversion we've talked about. 202 00:12:35,310 --> 00:12:39,390 Conditioned taste aversion is the usual phenomenon 203 00:12:39,390 --> 00:12:40,060 we talk about. 204 00:12:40,060 --> 00:12:43,300 Then conditioned action, conditioned appetitive behavior 205 00:12:43,300 --> 00:12:44,890 directed at quiescence, when there's 206 00:12:44,890 --> 00:12:52,700 some very strong annoyer like hunger or thirst, 207 00:12:52,700 --> 00:12:57,400 or anything that's extremely bothersome 208 00:12:57,400 --> 00:12:59,960 and difficult and uncomfortable. 209 00:12:59,960 --> 00:13:01,850 And finally operant conditioning, 210 00:13:01,850 --> 00:13:04,560 which is not stimulus conditioning at all 211 00:13:04,560 --> 00:13:08,940 but response selection, which Lorenz claims is much less 212 00:13:08,940 --> 00:13:15,600 common than people tend to be taught 213 00:13:15,600 --> 00:13:19,500 that learn about what they call instrumental conditioning. 214 00:13:19,500 --> 00:13:21,960 It's usually operant conditioning, 215 00:13:21,960 --> 00:13:25,430 which involves response selection. 216 00:13:25,430 --> 00:13:27,770 This is just an introduction to that. 217 00:13:27,770 --> 00:13:28,880 Lorenz. 218 00:13:28,880 --> 00:13:33,890 We've been talking mainly about Type S or stimulus selection 219 00:13:33,890 --> 00:13:36,830 types of learning. 220 00:13:36,830 --> 00:13:40,200 He says much exploratory behavior is that way, 221 00:13:40,200 --> 00:13:42,320 and it's the most common. 222 00:13:42,320 --> 00:13:44,700 Type R conditioning, response selection, 223 00:13:44,700 --> 00:13:48,170 is a little more rare. 224 00:13:48,170 --> 00:13:50,620 It includes operant conditioning, 225 00:13:50,620 --> 00:13:52,700 like with a confined animal that's 226 00:13:52,700 --> 00:13:55,195 trying every response he can in order to escape. 227 00:13:59,300 --> 00:14:02,540 It involves reinforcement, some kind of reward. 228 00:14:02,540 --> 00:14:05,136 He likes the word encouraging rather than reinforcement. 229 00:14:08,050 --> 00:14:10,540 And then he talks about, applies this 230 00:14:10,540 --> 00:14:12,740 to the study of fixed action patterns, where 231 00:14:12,740 --> 00:14:16,640 you have-- it starts with the drive, 232 00:14:16,640 --> 00:14:19,540 the action-specific potential which leads to appetitive 233 00:14:19,540 --> 00:14:25,020 behavior, which basically he's looking for the stimulus that 234 00:14:25,020 --> 00:14:31,190 can release the response of that response side of the fixed 235 00:14:31,190 --> 00:14:33,110 action pattern. 236 00:14:33,110 --> 00:14:35,740 So then his innate releasing mechanism 237 00:14:35,740 --> 00:14:40,215 is activated, which causes the consummatory behavior, 238 00:14:40,215 --> 00:14:42,410 the consummatory response. 239 00:14:42,410 --> 00:14:49,520 And then the feedback from that behavior, a the feedback 240 00:14:49,520 --> 00:14:52,560 affects the action-specific potential. 241 00:14:52,560 --> 00:14:53,430 It lowers it. 242 00:14:58,860 --> 00:15:02,740 It usually stops the consummatory behavior, 243 00:15:02,740 --> 00:15:03,735 reduces the drive. 244 00:15:07,880 --> 00:15:11,620 And so it reduces appetitive behavior. 245 00:15:11,620 --> 00:15:16,630 And it can actually shape the original releasing mechanism, 246 00:15:16,630 --> 00:15:21,350 because there's-- learning in these innate behavior sequences 247 00:15:21,350 --> 00:15:24,200 is mostly on the stimulus side. 248 00:15:24,200 --> 00:15:31,150 Like an animal that has built-in responses. 249 00:15:31,150 --> 00:15:34,600 We talked about the smiling response. 250 00:15:34,600 --> 00:15:36,570 It's a fixed action pattern. 251 00:15:36,570 --> 00:15:38,555 They will smile spontaneously sometimes, 252 00:15:38,555 --> 00:15:41,880 but they normally respond in almost reflex fashion 253 00:15:41,880 --> 00:15:44,650 to this specific configuration. 254 00:15:44,650 --> 00:15:48,390 And yet over time it acquires other characteristics, 255 00:15:48,390 --> 00:15:50,420 and his smiling becomes more limited, 256 00:15:50,420 --> 00:15:53,410 restricted to certain situations. 257 00:15:53,410 --> 00:15:54,570 OK 258 00:15:54,570 --> 00:15:56,440 And he has this interesting statement 259 00:15:56,440 --> 00:15:59,300 about why behaviorists have oversimplified 260 00:15:59,300 --> 00:16:01,820 the problem of learning and instinct. 261 00:16:01,820 --> 00:16:05,480 And he's talking about American comparative psychology. 262 00:16:05,480 --> 00:16:09,040 He says, they hope to be able to abstract generally valid laws 263 00:16:09,040 --> 00:16:12,780 prevailing in all learning processes, if not all behavior. 264 00:16:12,780 --> 00:16:16,420 And there have been many books written like this, almost all 265 00:16:16,420 --> 00:16:17,935 over here in America. 266 00:16:22,280 --> 00:16:24,650 In this way they hope to find a shortcut 267 00:16:24,650 --> 00:16:27,110 to an understanding of animal and human behavior 268 00:16:27,110 --> 00:16:30,740 without going to the trouble of analyzing the immensely 269 00:16:30,740 --> 00:16:33,090 complicated physiological machinery 270 00:16:33,090 --> 00:16:35,870 whose function is behavior. 271 00:16:35,870 --> 00:16:37,440 And I couldn't agree with him more. 272 00:16:41,370 --> 00:16:43,820 So these are the different kinds of learning affected 273 00:16:43,820 --> 00:16:47,260 by the consequences of behavior, the major categories. 274 00:16:47,260 --> 00:16:50,100 Start with conditioned appetitive behavior. 275 00:16:50,100 --> 00:16:54,720 And he says that that and conditioned appetitive behavior 276 00:16:54,720 --> 00:16:58,440 directed at quiescence are the most common types. 277 00:16:58,440 --> 00:17:02,020 And I also point out that usually operant conditioning 278 00:17:02,020 --> 00:17:04,079 is lumped together with conditioned appetitive 279 00:17:04,079 --> 00:17:06,050 behavior, even though they really 280 00:17:06,050 --> 00:17:09,250 are quite separable, as Lorenz points out. 281 00:17:11,880 --> 00:17:15,180 And as you know, sometimes-- like the Russians 282 00:17:15,180 --> 00:17:19,089 interpreted all behavior in terms of conditioned reflexes 283 00:17:19,089 --> 00:17:20,400 for many, many, many years. 284 00:17:23,680 --> 00:17:26,585 I don't know, people are natural lumpers, maybe. 285 00:17:26,585 --> 00:17:28,990 But Lorenz is more of a splitter here. 286 00:17:28,990 --> 00:17:30,980 He likes to divide it up because-- 287 00:17:30,980 --> 00:17:34,400 and I think this is much more relevant to any brain studies, 288 00:17:34,400 --> 00:17:37,190 because if you can't separate them, 289 00:17:37,190 --> 00:17:39,890 you won't be looking for different brain mechanisms. 290 00:17:39,890 --> 00:17:42,980 And in fact there are many distinct brain mechanisms 291 00:17:42,980 --> 00:17:47,480 going on, which we'll talk about at the end here. 292 00:17:47,480 --> 00:17:49,810 So these are examples of conditioned appetitive 293 00:17:49,810 --> 00:17:50,830 behavior. 294 00:17:50,830 --> 00:17:55,560 Von Frisch, Karl Von Frisch, had kept a pet fish. 295 00:17:55,560 --> 00:18:00,880 And he always whistled for it before he fed it. 296 00:18:00,880 --> 00:18:05,280 So the fish learned to begin searching for food 297 00:18:05,280 --> 00:18:06,760 as soon as he heard that whistle. 298 00:18:09,760 --> 00:18:13,650 The honey bees studied by Von Frisch. 299 00:18:13,650 --> 00:18:16,820 The insect searches for the stimulus situation 300 00:18:16,820 --> 00:18:18,690 which proved rewarding in the past. 301 00:18:18,690 --> 00:18:20,830 For example, you could teach it to fly 302 00:18:20,830 --> 00:18:24,190 to specifically colored patches. 303 00:18:24,190 --> 00:18:29,220 It's a type of learning that he's 304 00:18:29,220 --> 00:18:33,840 rewarded if he goes to those colors. 305 00:18:33,840 --> 00:18:37,350 Now the study of Pavlov's dogs. 306 00:18:37,350 --> 00:18:42,255 There's an interesting story about that from Howard Liddell. 307 00:18:48,580 --> 00:18:51,777 I can actually read what he said about that, 308 00:18:51,777 --> 00:18:53,110 because it's pretty interesting. 309 00:18:53,110 --> 00:18:53,985 He's quoting Liddell. 310 00:18:58,570 --> 00:19:03,172 In most of Pavlov's experiments, active appetitive behavior 311 00:19:03,172 --> 00:19:06,390 is made invisible by shackling the dog 312 00:19:06,390 --> 00:19:10,200 to a framework, the experimental apparatus, 313 00:19:10,200 --> 00:19:14,840 so that salivation is just about the only response which is not 314 00:19:14,840 --> 00:19:15,590 precluded. 315 00:19:15,590 --> 00:19:17,070 He can't do anything else. 316 00:19:17,070 --> 00:19:20,360 He's locked in the apparatus. 317 00:19:20,360 --> 00:19:24,500 And then Howard Liddell told him how he once conditioned the dog 318 00:19:24,500 --> 00:19:28,130 to salivate, using the conventional Pavlovian 319 00:19:28,130 --> 00:19:34,370 method, whenever a constantly ticking metronome was 320 00:19:34,370 --> 00:19:37,180 made to accelerate its beat. 321 00:19:37,180 --> 00:19:39,660 So he trained it, and he got the salivation 322 00:19:39,660 --> 00:19:42,190 in the typical Pavlovian sense. 323 00:19:45,910 --> 00:19:49,780 Then he untied the dog, and as soon 324 00:19:49,780 --> 00:19:52,480 as the dog was released from the apparatus, 325 00:19:52,480 --> 00:19:55,920 it ran up to the metronome at once, 326 00:19:55,920 --> 00:19:58,480 whined, wagged its tail violently, 327 00:19:58,480 --> 00:20:00,750 and pushed against the metronome with its nose, 328 00:20:00,750 --> 00:20:03,400 salivating intensely all the while, 329 00:20:03,400 --> 00:20:07,140 even though the metronome had not changed its rhythm. 330 00:20:07,140 --> 00:20:09,013 What had really been conditioned is 331 00:20:09,013 --> 00:20:12,010 that the dog was-- and the dog wasn't a reflex. 332 00:20:12,010 --> 00:20:15,060 There's no conditioned reflex here. 333 00:20:15,060 --> 00:20:18,470 But rather it was a complex and specific system 334 00:20:18,470 --> 00:20:22,729 of appetitive behavior by which a dog begs for food. 335 00:20:22,729 --> 00:20:23,895 That's what was conditioned. 336 00:20:27,610 --> 00:20:32,810 It's an interesting contrast, because what 337 00:20:32,810 --> 00:20:36,410 we call classical conditioning now in the laboratory, 338 00:20:36,410 --> 00:20:41,470 like conditioned eye blink in response to specific stimuli, 339 00:20:41,470 --> 00:20:43,360 is really very different from this. 340 00:20:46,100 --> 00:20:48,215 Anyway, this is conditioned appetitive behavior. 341 00:20:51,590 --> 00:20:54,680 He talks then about nest building in social weaver 342 00:20:54,680 --> 00:20:57,110 birds, or corvine birds. 343 00:20:57,110 --> 00:20:59,900 like crows and ravens. 344 00:20:59,900 --> 00:21:02,130 The weavers have an innate preference 345 00:21:02,130 --> 00:21:04,180 for a particular kind of grass. 346 00:21:04,180 --> 00:21:06,310 So if they get that grass, that's 347 00:21:06,310 --> 00:21:07,860 what they bring to the nest. 348 00:21:07,860 --> 00:21:10,540 But the corvines, like the jackdaws and the ravens, 349 00:21:10,540 --> 00:21:13,790 they use whatever nest material gives good feedback 350 00:21:13,790 --> 00:21:16,850 during nest-building actions. 351 00:21:16,850 --> 00:21:21,420 So they get twigs of the right shape and size, little twigs. 352 00:21:21,420 --> 00:21:24,700 He says it causes an orgastic climax when they're 353 00:21:24,700 --> 00:21:27,760 stuck in the nest because it gives them the right feedback. 354 00:21:27,760 --> 00:21:34,060 It gives them a real joy, and it rewards them. 355 00:21:34,060 --> 00:21:38,830 In fact, they can get supernormal feedback 356 00:21:38,830 --> 00:21:41,670 if they use soft metal wire because it gives 357 00:21:41,670 --> 00:21:43,885 such strong feedback, it works so well 358 00:21:43,885 --> 00:21:46,250 on that particular response. 359 00:21:46,250 --> 00:21:50,210 They become addicted to it, just like what 360 00:21:50,210 --> 00:21:53,800 happens in human addictions, even though the wire doesn't 361 00:21:53,800 --> 00:21:56,380 make a very good nest. 362 00:21:56,380 --> 00:21:59,890 But it gives the right feedback. 363 00:21:59,890 --> 00:22:02,800 Now when they evolved, of course, 364 00:22:02,800 --> 00:22:04,525 there wasn't wire mesh around. 365 00:22:08,010 --> 00:22:09,590 So there wasn't anything like that 366 00:22:09,590 --> 00:22:11,600 that could give that super normal feedback 367 00:22:11,600 --> 00:22:15,190 and result in maladaptive behavior. 368 00:22:15,190 --> 00:22:19,520 But because of humans, now they can 369 00:22:19,520 --> 00:22:26,890 acquire this kind of abnormal addiction to the wire. 370 00:22:26,890 --> 00:22:27,400 All right. 371 00:22:27,400 --> 00:22:32,100 So that's conditioned appetitive behavior. 372 00:22:32,100 --> 00:22:36,610 Now a little bit about conditioned aversion, 373 00:22:36,610 --> 00:22:38,530 the second kind of learning affected 374 00:22:38,530 --> 00:22:41,900 by the consequences of behavior. 375 00:22:41,900 --> 00:22:45,490 This is John Garcia's discovery that we've talked about, 376 00:22:45,490 --> 00:22:47,420 the poison bait effect. 377 00:22:47,420 --> 00:22:53,070 I just want to point out that the sickness, the response 378 00:22:53,070 --> 00:23:01,040 here, the nausea, the illness, that becomes 379 00:23:01,040 --> 00:23:04,650 associated with a stimulus, not the most recently 380 00:23:04,650 --> 00:23:08,050 encountered stimulus. 381 00:23:08,050 --> 00:23:12,550 It associates the most novel previous stimulus. 382 00:23:12,550 --> 00:23:16,780 Now I'd like to know, are there other cases 383 00:23:16,780 --> 00:23:19,280 of conditioned aversion other than 384 00:23:19,280 --> 00:23:22,810 ingestion-related behaviors? 385 00:23:22,810 --> 00:23:24,840 It's likely that there are, but this 386 00:23:24,840 --> 00:23:29,000 is the one that's been most studied. 387 00:23:29,000 --> 00:23:30,790 Lorenz believes that conditioned aversion 388 00:23:30,790 --> 00:23:33,990 is more common in many animals in nature 389 00:23:33,990 --> 00:23:36,870 than is something like operant conditioning, which 390 00:23:36,870 --> 00:23:38,290 we will talk about in a minute. 391 00:23:41,890 --> 00:23:45,500 And I just remind you that Garcia's discovery of this 392 00:23:45,500 --> 00:23:49,190 was initially suppressed by the editors of American behavior 393 00:23:49,190 --> 00:23:50,260 magazines. 394 00:23:50,260 --> 00:23:56,960 It didn't fit the beliefs about conditioning and learning. 395 00:24:00,100 --> 00:24:02,920 They're not supposed to become conditioned to something that 396 00:24:02,920 --> 00:24:05,990 occurred a lot earlier, but only the most recent stimuli. 397 00:24:08,930 --> 00:24:13,130 But actually they become conditioned to the more novel 398 00:24:13,130 --> 00:24:21,620 stimulus, even though it might have occurred much earlier. 399 00:24:21,620 --> 00:24:25,620 And then conditioned action. 400 00:24:25,620 --> 00:24:28,970 This is usually an artificial kind of motor learning, 401 00:24:28,970 --> 00:24:32,830 like you shape actions by circus animals. 402 00:24:32,830 --> 00:24:37,090 You-- I said initially elicit a fixed action 403 00:24:37,090 --> 00:24:39,110 pattern in response to a command. 404 00:24:39,110 --> 00:24:44,230 You're actually eliciting a fixed motor pattern in response 405 00:24:44,230 --> 00:24:45,000 to a command. 406 00:24:45,000 --> 00:24:50,290 So the horse's capriole, that is the leap and kicking 407 00:24:50,290 --> 00:24:56,130 that a horse does to try to get rid of a predator that's 408 00:24:56,130 --> 00:25:00,570 attacking it-- they will learn to perform that just 409 00:25:00,570 --> 00:25:01,370 to get sugar. 410 00:25:01,370 --> 00:25:03,105 So that's a conditioned action. 411 00:25:05,980 --> 00:25:08,360 Von Frisch conditioned a parakeet 412 00:25:08,360 --> 00:25:11,620 to defecate in order to be released from his stage. 413 00:25:11,620 --> 00:25:14,590 The reward was getting released from his cage. 414 00:25:14,590 --> 00:25:16,985 So that was certainly a conditioned action, 415 00:25:16,985 --> 00:25:19,870 a kind of artificial learning. 416 00:25:19,870 --> 00:25:23,000 But then he points out that a lot of actions you just 417 00:25:23,000 --> 00:25:23,900 can't do that with. 418 00:25:23,900 --> 00:25:25,550 You can't condition them like that. 419 00:25:25,550 --> 00:25:30,260 Like tendon reflexes, sexual action 420 00:25:30,260 --> 00:25:31,670 patterns in most animals. 421 00:25:31,670 --> 00:25:34,270 You can't condition like that. 422 00:25:34,270 --> 00:25:36,190 Bill shaking in mallards. 423 00:25:36,190 --> 00:25:38,910 And I did a lot of work on hamsters. 424 00:25:38,910 --> 00:25:41,095 And I tried to condition their orienting. 425 00:25:45,950 --> 00:25:50,281 For example, normally you present the stimulus here, 426 00:25:50,281 --> 00:25:50,780 he turns. 427 00:25:50,780 --> 00:25:52,470 It's an innate response. 428 00:25:52,470 --> 00:25:55,080 And we know the pathway in the brain from retina 429 00:25:55,080 --> 00:25:57,830 to the midbrain tectum and the superior colliculus 430 00:25:57,830 --> 00:25:58,690 that controls it. 431 00:25:58,690 --> 00:26:02,950 And then we know the output pathway as well. 432 00:26:02,950 --> 00:26:06,110 But what if you never rewarded them when they turn? 433 00:26:06,110 --> 00:26:11,816 You reward them over here and stimulate their whiskers 434 00:26:11,816 --> 00:26:12,315 afterwards. 435 00:26:12,315 --> 00:26:15,300 So you present it visually over here. 436 00:26:15,300 --> 00:26:17,230 And then you immediately touch the whiskers 437 00:26:17,230 --> 00:26:19,320 and they turn and get the reward. 438 00:26:19,320 --> 00:26:20,470 What happens? 439 00:26:20,470 --> 00:26:25,180 Well, you've seen the conditioning action. 440 00:26:25,180 --> 00:26:27,100 He does learn. 441 00:26:27,100 --> 00:26:29,340 But if you look at the behavior in detail, 442 00:26:29,340 --> 00:26:33,050 you'll see that actually there's a little hesitation. 443 00:26:33,050 --> 00:26:35,560 He'll usually start to turn the correct direction, 444 00:26:35,560 --> 00:26:39,080 and then he'll inhibit that. 445 00:26:39,080 --> 00:26:41,780 He learns to inhibit the behavior 446 00:26:41,780 --> 00:26:44,040 and then turn to get the reward. 447 00:26:44,040 --> 00:26:47,860 And if he's really learned that and there's 448 00:26:47,860 --> 00:26:50,650 any disturbance, like a novel stimulus, something 449 00:26:50,650 --> 00:26:56,100 that changes the environment a little bit, 450 00:26:56,100 --> 00:26:59,940 the normal behavior comes right back. 451 00:26:59,940 --> 00:27:02,160 You can't release press. 452 00:27:02,160 --> 00:27:07,930 You can't condition it in the sense of the earlier examples 453 00:27:07,930 --> 00:27:09,580 here. 454 00:27:09,580 --> 00:27:11,290 The innate responses will dominate. 455 00:27:16,180 --> 00:27:18,120 The other situation was when I was 456 00:27:18,120 --> 00:27:21,620 able to get the optic tract to literally go 457 00:27:21,620 --> 00:27:25,020 to the wrong side of the brain by an early brain lesion. 458 00:27:25,020 --> 00:27:27,139 So again, you present the stimulus, 459 00:27:27,139 --> 00:27:28,680 they always turn the wrong direction. 460 00:27:28,680 --> 00:27:31,290 But they never get rewarded for that 461 00:27:31,290 --> 00:27:33,230 unless you do it artificially. 462 00:27:33,230 --> 00:27:35,030 So what happens? 463 00:27:35,030 --> 00:27:40,010 I found that in that area of the visual field that's affected, 464 00:27:40,010 --> 00:27:43,510 they never do learn, in spite of them always being rewarded. 465 00:27:47,749 --> 00:27:49,040 Because they're never rewarded. 466 00:27:49,040 --> 00:27:52,430 They never learn to turn in the right direction 467 00:27:52,430 --> 00:27:55,590 even though it would be adaptive to do so, 468 00:27:55,590 --> 00:27:59,150 because the connection won't change just 469 00:27:59,150 --> 00:28:01,580 because of learning. 470 00:28:01,580 --> 00:28:04,555 So that's what's underlying the innate behavior. 471 00:28:07,250 --> 00:28:12,420 And even if he learns to totally suppress, 472 00:28:12,420 --> 00:28:17,550 that you will bring out-- it's very easy to train him. 473 00:28:17,550 --> 00:28:22,350 One reward where you're getting the wrong direction training 474 00:28:22,350 --> 00:28:24,460 and it all comes right back. 475 00:28:24,460 --> 00:28:26,722 So conditioned appetitive behavior 476 00:28:26,722 --> 00:28:27,680 directed at quiescence. 477 00:28:27,680 --> 00:28:30,050 Now this is very common. 478 00:28:30,050 --> 00:28:35,580 We're talking here about annoyers, tension reduction. 479 00:28:35,580 --> 00:28:40,215 The tension or the annoyer is hunger or thirst or stress, 480 00:28:40,215 --> 00:28:44,040 a state of stress or conflict. 481 00:28:44,040 --> 00:28:48,530 You classify habitat selection this way 482 00:28:48,530 --> 00:28:51,400 because when an animal's searching for a habitat, 483 00:28:51,400 --> 00:28:54,290 it's another drive that's not being satisfied. 484 00:28:54,290 --> 00:28:55,180 He's uncomfortable. 485 00:28:55,180 --> 00:28:58,020 The drive is high, and it can only be satisfied 486 00:28:58,020 --> 00:29:01,450 if he finds a satisfactory habitat. 487 00:29:01,450 --> 00:29:07,490 And that reduces the tension and causes his reward. 488 00:29:07,490 --> 00:29:09,910 So that's conditioned appetitive behavior 489 00:29:09,910 --> 00:29:11,880 directed at quiescence. 490 00:29:11,880 --> 00:29:15,560 So it usually involves a search for a stimulus situation 491 00:29:15,560 --> 00:29:16,770 that leads to a reward. 492 00:29:16,770 --> 00:29:20,400 It's another kind of stimulus selection. 493 00:29:20,400 --> 00:29:24,280 The reward is the reduction in the tension, the stress, 494 00:29:24,280 --> 00:29:28,970 the anxiety, the annoyance, whatever it is. 495 00:29:28,970 --> 00:29:34,920 So most of the kind of learning studied in rats, 496 00:29:34,920 --> 00:29:37,980 where they're looking at what we call instrumental conditioning, 497 00:29:37,980 --> 00:29:41,350 is actually stimulus selection. 498 00:29:41,350 --> 00:29:46,110 So we'll talk about response selection. 499 00:29:46,110 --> 00:29:50,120 Now, operant conditioning. 500 00:29:50,120 --> 00:29:51,385 Very different from this. 501 00:29:54,550 --> 00:29:58,750 Even though a lot of scientists group these two, conditioned 502 00:29:58,750 --> 00:30:01,360 appetitive behavior directed at quiescence and operant 503 00:30:01,360 --> 00:30:03,640 conditioning, together. 504 00:30:03,640 --> 00:30:06,270 So this is a kind of response selection. 505 00:30:06,270 --> 00:30:09,570 Very highly studied, like in Skinnerian conditioning. 506 00:30:09,570 --> 00:30:12,540 Like you put a cat in a puzzle box, 507 00:30:12,540 --> 00:30:17,960 and he will try every response that he's able. 508 00:30:17,960 --> 00:30:19,960 He will try everything in order to-- 509 00:30:19,960 --> 00:30:23,380 and eventually, if possible, he'll 510 00:30:23,380 --> 00:30:25,080 discover some response that helps 511 00:30:25,080 --> 00:30:27,310 him get out of the puzzle box. 512 00:30:27,310 --> 00:30:33,050 You put a male dog eager to mate, 513 00:30:33,050 --> 00:30:37,760 and the female dog, the bitch, is confined, 514 00:30:37,760 --> 00:30:39,670 but there is some way to get to her, 515 00:30:39,670 --> 00:30:44,210 he will try every response in order to get to her. 516 00:30:44,210 --> 00:30:45,700 So that's response selection. 517 00:30:45,700 --> 00:30:49,820 He's trying to find a response that works. 518 00:30:49,820 --> 00:30:55,040 He points out that in nature operant conditioning is rarer 519 00:30:55,040 --> 00:30:58,190 than generally assumed. 520 00:30:58,190 --> 00:31:00,970 It occurs in situations of appetitive behavior directed 521 00:31:00,970 --> 00:31:03,360 at quiescence, or in exploratory behavior. 522 00:31:06,460 --> 00:31:08,885 Now we come to real motor learning, 523 00:31:08,885 --> 00:31:13,510 where the motor responses themselves can change. 524 00:31:16,560 --> 00:31:19,086 We know that skilled movements can be shaped. 525 00:31:23,350 --> 00:31:27,710 We try this in sports all the time. 526 00:31:27,710 --> 00:31:32,070 In nature you'll see animals acquire path habits. 527 00:31:32,070 --> 00:31:34,420 When they acquire a path habit they 528 00:31:34,420 --> 00:31:36,905 can follow that path with much greater speed. 529 00:31:40,300 --> 00:31:42,860 It includes things like recitation from memory. 530 00:31:42,860 --> 00:31:45,360 You can learn to rattle off a poem. 531 00:31:45,360 --> 00:31:47,690 Even if you forget even what the poem's about, 532 00:31:47,690 --> 00:31:51,290 you can learn all the words and everything. 533 00:31:51,290 --> 00:31:53,200 And there are examples in animal behavior. 534 00:31:53,200 --> 00:31:57,920 For example, a little vole kept as a pet by, 535 00:31:57,920 --> 00:32:02,340 I think it was Lorenz, always followed, 536 00:32:02,340 --> 00:32:06,450 ran in a certain pattern to get back to his nest. 537 00:32:06,450 --> 00:32:08,244 And animals learn this all the time. 538 00:32:08,244 --> 00:32:09,910 When they're in their local environment, 539 00:32:09,910 --> 00:32:14,600 they learn actions that bring them rapidly back to the nest. 540 00:32:14,600 --> 00:32:17,380 And if there's one action that they are commonly 541 00:32:17,380 --> 00:32:21,170 done-- like it might involve leaping 542 00:32:21,170 --> 00:32:24,760 onto an obstacle on their way to the nest. 543 00:32:24,760 --> 00:32:28,210 Well, suddenly you just take the obstacle away. 544 00:32:28,210 --> 00:32:32,610 This is how you show it's a learned motor pattern. 545 00:32:32,610 --> 00:32:34,470 He has shaped his motor behavior, 546 00:32:34,470 --> 00:32:36,460 because he still goes through the whole thing. 547 00:32:36,460 --> 00:32:39,870 And suddenly that object's not there to land on. 548 00:32:39,870 --> 00:32:40,750 And what happens? 549 00:32:40,750 --> 00:32:45,600 He leaps anyway and hits a big surprise, because he falls 550 00:32:45,600 --> 00:32:49,140 further and lands on the ground. 551 00:32:49,140 --> 00:32:52,200 These kinds of learned habits can 552 00:32:52,200 --> 00:32:54,650 be pretty resistant to change. 553 00:32:54,650 --> 00:32:57,330 They have a lot of similarities to fixed motor patterns. 554 00:32:57,330 --> 00:33:01,220 In fact, they acquire their own appetitive behavior, 555 00:33:01,220 --> 00:33:01,955 in many cases. 556 00:33:05,020 --> 00:33:08,140 People that learn particular patterns of movement in sports 557 00:33:08,140 --> 00:33:10,440 can become very motivated to do those things. 558 00:33:14,780 --> 00:33:17,860 And we know that different parts of the brain 559 00:33:17,860 --> 00:33:23,370 are involved in changing motor patterns, especially 560 00:33:23,370 --> 00:33:24,925 cerebellar mechanisms. 561 00:33:31,070 --> 00:33:34,340 Just more about shaping of motor patterns. 562 00:33:34,340 --> 00:33:37,650 There's a lot of species differences here. 563 00:33:37,650 --> 00:33:42,250 Let's just remind ourselves of species differences 564 00:33:42,250 --> 00:33:47,560 in locomotion, because it gives an example of what you can 565 00:33:47,560 --> 00:33:50,970 train and what you cannot train an animal to do. 566 00:33:50,970 --> 00:33:54,450 So horses are very adapted to running on flat ground. 567 00:33:54,450 --> 00:33:57,210 They can't adjust well to uneven terrain 568 00:33:57,210 --> 00:33:59,270 no matter how much you try to train them. 569 00:33:59,270 --> 00:34:04,320 They're very limited in the responses available to them, 570 00:34:04,320 --> 00:34:06,530 the types of movements they can make. 571 00:34:06,530 --> 00:34:08,500 Donkeys are more sure-footed because they've 572 00:34:08,500 --> 00:34:10,639 got more responses available. 573 00:34:10,639 --> 00:34:15,415 Goats, the chamois, they're adapted to hills and mountains. 574 00:34:15,415 --> 00:34:16,719 They're very sure-footed. 575 00:34:16,719 --> 00:34:19,159 So what is that sure-footedness? 576 00:34:19,159 --> 00:34:23,110 Smaller units of action. 577 00:34:23,110 --> 00:34:31,980 And with smaller units, they can adjust direction, 578 00:34:31,980 --> 00:34:38,330 adjust for foot placement more readily in these rapid actions. 579 00:34:38,330 --> 00:34:41,780 In other words, higher control has more possibilities. 580 00:34:41,780 --> 00:34:45,030 There's more movement patterns to control. 581 00:34:45,030 --> 00:34:46,960 And that leads to a discussion of what 582 00:34:46,960 --> 00:34:49,830 we mean by voluntary action. 583 00:34:49,830 --> 00:34:53,065 He points out that arboreal creatures, 584 00:34:53,065 --> 00:34:59,360 they need a lot more action patterns 585 00:34:59,360 --> 00:35:00,965 to negotiate the trees. 586 00:35:06,030 --> 00:35:10,000 So it's a lot easier to shape movements 587 00:35:10,000 --> 00:35:13,000 in primates and squirrels and tree shrews 588 00:35:13,000 --> 00:35:14,780 and so forth that live in trees. 589 00:35:18,780 --> 00:35:22,140 We talk about will or volition when 590 00:35:22,140 --> 00:35:25,340 it really amounts to higher level initiation of fixed motor 591 00:35:25,340 --> 00:35:30,050 patterns, or learned sequences, skilled movements. 592 00:35:30,050 --> 00:35:32,930 And we've already talked about species differences 593 00:35:32,930 --> 00:35:37,050 on how small the elements of movements are. 594 00:35:37,050 --> 00:35:40,950 And the example he gives about voluntary movement and insight 595 00:35:40,950 --> 00:35:46,110 concerns his studies of geese. 596 00:35:46,110 --> 00:35:50,020 He says that their spatial insight can exceed their motor 597 00:35:50,020 --> 00:35:54,050 abilities because they don't have so many different motor 598 00:35:54,050 --> 00:35:55,310 patterns available to them. 599 00:35:55,310 --> 00:35:58,520 So you can teach them to climb stairs fairly well. 600 00:35:58,520 --> 00:36:01,000 But when you try to teach them how to descend a stairway, 601 00:36:01,000 --> 00:36:05,290 they can't even learn to adjust the length of their stride 602 00:36:05,290 --> 00:36:05,790 well enough. 603 00:36:08,950 --> 00:36:10,910 And they will stumble. 604 00:36:10,910 --> 00:36:14,065 They have a lot of trouble going down the stairs. 605 00:36:16,840 --> 00:36:20,550 So species differ a lot in how much 606 00:36:20,550 --> 00:36:25,520 they can control when they want to. 607 00:36:25,520 --> 00:36:27,390 Voluntary control, we call it. 608 00:36:27,390 --> 00:36:31,120 It's the influence of motivational systems 609 00:36:31,120 --> 00:36:35,080 on higher level control systems. 610 00:36:35,080 --> 00:36:36,800 And then he comes to a discussion 611 00:36:36,800 --> 00:36:38,090 of exploratory behavior. 612 00:36:48,800 --> 00:36:50,550 Exploratory behavior or curiosity. 613 00:36:53,794 --> 00:36:55,085 And this is pretty interesting. 614 00:36:58,820 --> 00:37:01,880 When you study exploratory behavior, of course, 615 00:37:01,880 --> 00:37:04,000 it's very influenced by novelty. 616 00:37:04,000 --> 00:37:05,690 Many animals are, in fact, highly 617 00:37:05,690 --> 00:37:09,570 motivated to explore novel situations, 618 00:37:09,570 --> 00:37:14,770 and they will direct multiple different patterns of behavior, 619 00:37:14,770 --> 00:37:16,170 sometimes unrelated. 620 00:37:16,170 --> 00:37:17,290 Even the little hamster. 621 00:37:17,290 --> 00:37:20,500 You put them in a novel room with different objects 622 00:37:20,500 --> 00:37:24,300 that they're not accustomed to seeing. 623 00:37:24,300 --> 00:37:25,460 It's amazing. 624 00:37:25,460 --> 00:37:30,960 Their initial response, if it's very novel, will be fright. 625 00:37:30,960 --> 00:37:33,520 Their tension levels go way up. 626 00:37:33,520 --> 00:37:37,200 But in fact, if you've got them in a situation 627 00:37:37,200 --> 00:37:39,206 that they're used to, you test them 628 00:37:39,206 --> 00:37:42,370 in an apparatus they're very familiar with, 629 00:37:42,370 --> 00:37:43,830 they will explore. 630 00:37:43,830 --> 00:37:48,750 And you will see them trying all different kinds 631 00:37:48,750 --> 00:37:49,500 of motor patterns. 632 00:37:49,500 --> 00:37:52,000 You'll see rapid switching between different behavior 633 00:37:52,000 --> 00:37:53,395 patterns or motor patterns. 634 00:37:58,610 --> 00:38:02,510 And when there's a strong action-specific potential 635 00:38:02,510 --> 00:38:06,130 behind that's causing those motor patterns, 636 00:38:06,130 --> 00:38:09,580 you don't see that switching from one to the other. 637 00:38:09,580 --> 00:38:13,050 So in exploratory behavior, there's 638 00:38:13,050 --> 00:38:21,030 a difference in the structure of what's behind the movement. 639 00:38:21,030 --> 00:38:24,350 And exploratory behavior is not as well understood 640 00:38:24,350 --> 00:38:28,600 as fixed action patterns, because now the motor part 641 00:38:28,600 --> 00:38:32,710 can become uncoupled with the normal action. 642 00:38:32,710 --> 00:38:34,320 The motivation here is curiosity. 643 00:38:37,020 --> 00:38:40,150 Very similar to play behavior. 644 00:38:40,150 --> 00:38:44,600 He says it occurs in a field devoid of tension. 645 00:38:44,600 --> 00:38:46,660 And yet the motivation to explore 646 00:38:46,660 --> 00:38:50,730 can be so strong, an animal sometimes prefer 647 00:38:50,730 --> 00:38:53,740 that rather than eating, even if he's hungry. 648 00:38:56,430 --> 00:39:00,340 And it does have a function, very important functions. 649 00:39:00,340 --> 00:39:02,835 When the environment changes, it helps the animal, 650 00:39:02,835 --> 00:39:07,890 it's adaptive for the animal to explore it. 651 00:39:07,890 --> 00:39:10,570 He needs to learn the spatial layout of the environment 652 00:39:10,570 --> 00:39:13,910 around him in relation to visual landmarks 653 00:39:13,910 --> 00:39:16,490 and other sensory cues, because if they change, 654 00:39:16,490 --> 00:39:19,200 then he's in trouble if he gets mixed up trying to get home, 655 00:39:19,200 --> 00:39:21,830 trying to escape. 656 00:39:21,830 --> 00:39:23,735 It makes his life a lot more efficient 657 00:39:23,735 --> 00:39:27,090 if he stays familiar with the novel situation. 658 00:39:27,090 --> 00:39:29,930 There were experiments on this using hamsters 659 00:39:29,930 --> 00:39:33,660 because they're so curious. 660 00:39:33,660 --> 00:39:35,480 Actually, my first publication's called 661 00:39:35,480 --> 00:39:37,330 "Curiosity and the Hamster." 662 00:39:37,330 --> 00:39:41,130 I was using exploratory behavior to reward them 663 00:39:41,130 --> 00:39:43,160 for doing other things. 664 00:39:43,160 --> 00:39:45,470 Reward them for pressing a bar, reward them 665 00:39:45,470 --> 00:39:47,380 for going through doors. 666 00:39:47,380 --> 00:39:49,750 It was so novel at the time, people still 667 00:39:49,750 --> 00:39:53,015 believed that this wasn't supposed to be possible. 668 00:39:55,740 --> 00:39:58,005 I had to play around at home with a pet hamster, 669 00:39:58,005 --> 00:40:01,190 and I found out I could teach him to press a bar made out 670 00:40:01,190 --> 00:40:04,604 of a knife taped to a baby rattle. 671 00:40:04,604 --> 00:40:06,020 And I could teach them to do this. 672 00:40:06,020 --> 00:40:08,900 And my professor was so intrigued by that, 673 00:40:08,900 --> 00:40:11,600 he got me to write a paper about it. 674 00:40:11,600 --> 00:40:13,140 And that was because of the theories 675 00:40:13,140 --> 00:40:21,030 that were dominating American psychology at that time. 676 00:40:21,030 --> 00:40:24,680 But Catherine Blanc in Europe-- I think it was in France, 677 00:40:24,680 --> 00:40:27,370 but I don't remember for sure anymore-- 678 00:40:27,370 --> 00:40:29,550 but she studied this experimentally. 679 00:40:29,550 --> 00:40:35,830 And she found out that in their exploratory behavior, 680 00:40:35,830 --> 00:40:37,730 they are acquiring knowledge that they 681 00:40:37,730 --> 00:40:40,070 can use in other situations. 682 00:40:40,070 --> 00:40:42,590 They become more efficient in finding their way 683 00:40:42,590 --> 00:40:43,465 in another situation. 684 00:40:46,030 --> 00:40:49,040 And then Lorenz points out that exploratory behavior 685 00:40:49,040 --> 00:40:55,550 is especially highly developed in unspecialized species. 686 00:40:55,550 --> 00:41:01,130 He says their specialization is being versatile, like humans, 687 00:41:01,130 --> 00:41:03,650 like rats, like ravens. 688 00:41:03,650 --> 00:41:06,350 These are animals that aren't so limited 689 00:41:06,350 --> 00:41:08,960 to one type of environment. 690 00:41:08,960 --> 00:41:12,900 We should add crows to that. 691 00:41:12,900 --> 00:41:22,010 Mice are certainly more general than most voles, for example, 692 00:41:22,010 --> 00:41:25,900 because they can adapt to more different situations. 693 00:41:25,900 --> 00:41:28,880 And all of this type of animal shows 694 00:41:28,880 --> 00:41:30,360 a lot of exploratory behavior. 695 00:41:35,080 --> 00:41:38,426 AUDIENCE: What would be a specialized species? 696 00:41:38,426 --> 00:41:41,060 PROFESSOR: Most of the species we've been talking about 697 00:41:41,060 --> 00:41:43,830 are pretty specialized. 698 00:41:43,830 --> 00:41:50,300 Every species will show some-- I think all of the mammals 699 00:41:50,300 --> 00:41:52,640 show some curiosity. 700 00:41:52,640 --> 00:41:55,930 But the amount that they will show will vary. 701 00:42:01,460 --> 00:42:04,460 What is characteristic about ravens, rats, and humans? 702 00:42:08,320 --> 00:42:10,190 The first thing you think of is that they've 703 00:42:10,190 --> 00:42:13,320 adapted to so many different environments, 704 00:42:13,320 --> 00:42:15,710 especially if you group the corvids together, 705 00:42:15,710 --> 00:42:16,925 the ravens and the crows. 706 00:42:20,950 --> 00:42:25,530 We know that rats can be found everywhere. 707 00:42:25,530 --> 00:42:28,420 The same is true for different corvids. 708 00:42:28,420 --> 00:42:31,690 They have specific adaptations, of course, 709 00:42:31,690 --> 00:42:36,670 that makes the jackdaw common in parts of Europe 710 00:42:36,670 --> 00:42:40,120 and the crow much more common over here. 711 00:42:40,120 --> 00:42:43,630 Humans, of course, are the most versatile, 712 00:42:43,630 --> 00:42:45,820 the least specialized. 713 00:42:45,820 --> 00:42:51,480 That doesn't mean-- we do depend a lot on learning. 714 00:42:51,480 --> 00:42:54,030 But what is different about-- for example, 715 00:42:54,030 --> 00:42:57,210 back here, when we're talking about shaping 716 00:42:57,210 --> 00:43:02,750 of motor learning here. 717 00:43:02,750 --> 00:43:08,360 It depends a lot on how many fixed action patterns you have. 718 00:43:08,360 --> 00:43:12,740 The more fixed action patterns that make up your movements, 719 00:43:12,740 --> 00:43:16,140 the more versatile you are, which is one reason humans 720 00:43:16,140 --> 00:43:17,980 are so versatile. 721 00:43:17,980 --> 00:43:21,165 We actually have more fixed action patterns than animals. 722 00:43:23,700 --> 00:43:26,875 At least on the motor side, we have many more choices. 723 00:43:38,840 --> 00:43:40,700 Some of the most specialized animals 724 00:43:40,700 --> 00:43:42,740 are the ones that are specialized for feeding 725 00:43:42,740 --> 00:43:46,820 on very particular things, like the koala. 726 00:43:46,820 --> 00:43:49,060 They only can eat certain things, 727 00:43:49,060 --> 00:43:52,260 and they don't survive well in other situations. 728 00:43:52,260 --> 00:43:55,530 Why aren't hamsters-- why haven't they 729 00:43:55,530 --> 00:43:58,320 spread all over the US? 730 00:43:58,320 --> 00:44:05,530 They're just not adaptable like the rat, or even the mouse. 731 00:44:05,530 --> 00:44:09,620 So they're only found-- they're native to Syria 732 00:44:09,620 --> 00:44:14,080 and the northern part of what is now Israel. 733 00:44:14,080 --> 00:44:18,560 And if you even move towards Turkey and that direction, 734 00:44:18,560 --> 00:44:19,730 the species changes. 735 00:44:19,730 --> 00:44:21,320 It's no longer Syrian hamster. 736 00:44:21,320 --> 00:44:24,730 It becomes a different hamster that has adapted specifically 737 00:44:24,730 --> 00:44:26,480 to the higher elevations. 738 00:44:26,480 --> 00:44:35,330 If you go north, northeast, you'll see the-- no, 739 00:44:35,330 --> 00:44:39,320 sorry, if you go further, you go, say, into Romania, again, 740 00:44:39,320 --> 00:44:41,160 another species appears. 741 00:44:41,160 --> 00:44:45,430 You go the other way, you start to get the Chinese hamster, 742 00:44:45,430 --> 00:44:47,890 or Siberian hamster if you go further. 743 00:44:47,890 --> 00:44:49,440 They're all different. 744 00:44:49,440 --> 00:44:50,910 They've adapted differently. 745 00:44:50,910 --> 00:44:53,069 But the rat, you keep finding the rat 746 00:44:53,069 --> 00:44:54,485 in all these different situations. 747 00:44:57,460 --> 00:44:59,790 So any species that's very limited to one 748 00:44:59,790 --> 00:45:05,080 habitat you wouldn't call a generalized species. 749 00:45:05,080 --> 00:45:09,400 And that's most of the ones that I'm talking about. 750 00:45:09,400 --> 00:45:09,900 OK. 751 00:45:09,900 --> 00:45:11,450 What about play behavior? 752 00:45:11,450 --> 00:45:15,740 Similar to explorative behavior. 753 00:45:15,740 --> 00:45:18,710 The very primitive play are simply-- 754 00:45:18,710 --> 00:45:21,530 you could call them in vacua reactions motivated 755 00:45:21,530 --> 00:45:26,880 by action-specific potentials or drives that are high. 756 00:45:26,880 --> 00:45:28,170 But there is a difference. 757 00:45:28,170 --> 00:45:35,540 You don't call it play if they execute the entire sequence 758 00:45:35,540 --> 00:45:41,440 from a high action-specific potential, a high drive level, 759 00:45:41,440 --> 00:45:45,040 to the motor pattern, like running and chasing 760 00:45:45,040 --> 00:45:47,610 playful fighting or prey catching. 761 00:45:47,610 --> 00:45:50,470 The animals will enjoy the motor patterns, 762 00:45:50,470 --> 00:45:53,080 and they will do this even before they're using them 763 00:45:53,080 --> 00:45:56,840 to catch prey, to feed and so forth. 764 00:45:56,840 --> 00:46:00,340 So it's very common in young animals. 765 00:46:00,340 --> 00:46:03,470 And he points out that play can shift 766 00:46:03,470 --> 00:46:05,990 to the real thing in some situations. 767 00:46:05,990 --> 00:46:07,610 There's a lot of danger in playing 768 00:46:07,610 --> 00:46:11,550 with an adult tomcat or badger. 769 00:46:11,550 --> 00:46:14,030 You might have tamed it and think it's your pet, 770 00:46:14,030 --> 00:46:22,900 but it can switch suddenly to the real thing 771 00:46:22,900 --> 00:46:24,840 when their motivation becomes high. 772 00:46:27,420 --> 00:46:29,630 So if you play with an adult tomcat, 773 00:46:29,630 --> 00:46:32,497 you have to be pretty careful because of that. 774 00:46:32,497 --> 00:46:33,580 You don't want to trigger. 775 00:46:33,580 --> 00:46:35,100 And that's certainly true. 776 00:46:35,100 --> 00:46:37,290 People get caught off guard all the time 777 00:46:37,290 --> 00:46:42,110 when they raise a tiger or other big cat in their home. 778 00:46:42,110 --> 00:46:47,310 Or even a champ or a monkey, because even though they 779 00:46:47,310 --> 00:46:49,650 seem to be playing, it can switch suddenly 780 00:46:49,650 --> 00:46:51,840 to the real thing. 781 00:46:51,840 --> 00:46:56,590 And they can kill you or your child or whatever. 782 00:46:56,590 --> 00:46:58,230 So it's dangerous to do that. 783 00:47:04,310 --> 00:47:08,335 He points out that, on the motivational side, 784 00:47:08,335 --> 00:47:13,310 a cat that has no opportunity to catch and kill prey 785 00:47:13,310 --> 00:47:16,970 can compensate by playing. 786 00:47:16,970 --> 00:47:19,190 It doesn't mean that it's always totally safe, 787 00:47:19,190 --> 00:47:20,774 but they appear to do that. 788 00:47:20,774 --> 00:47:22,190 So how do you tell the difference? 789 00:47:22,190 --> 00:47:25,530 Well, one way is physiologically. 790 00:47:25,530 --> 00:47:28,890 Because in play, the autonomic nervous system 791 00:47:28,890 --> 00:47:30,840 isn't involved in the same way. 792 00:47:30,840 --> 00:47:33,030 So if you major autonomic responses, 793 00:47:33,030 --> 00:47:38,730 you'll find big differences between the way the heart rate, 794 00:47:38,730 --> 00:47:40,430 the breathing, blood pressure changes, 795 00:47:40,430 --> 00:47:42,530 and so forth are responding in the two situations. 796 00:47:45,062 --> 00:47:46,520 Now what are the functions of play? 797 00:47:46,520 --> 00:47:51,010 And I just felt-- in the last half of his career studied 798 00:47:51,010 --> 00:47:54,270 humans and wrote the book Human Ecology. 799 00:47:54,270 --> 00:47:59,600 He studied development of play in cats and other carnivores. 800 00:47:59,600 --> 00:48:03,970 He found evidence of their learning of coordination, 801 00:48:03,970 --> 00:48:08,380 learning of stimulus selection, so they could change, 802 00:48:08,380 --> 00:48:11,605 so they would respond to more relevant stimuli. 803 00:48:14,260 --> 00:48:17,150 And also they would invent, in their play 804 00:48:17,150 --> 00:48:19,850 they would invent movement patterns simply 805 00:48:19,850 --> 00:48:25,940 by linking different elements, the different inherited 806 00:48:25,940 --> 00:48:29,070 elements of their behavior. 807 00:48:29,070 --> 00:48:31,120 And they will do that even though it 808 00:48:31,120 --> 00:48:32,530 doesn't lead to any reward. 809 00:48:32,530 --> 00:48:38,050 The reward is simply doing it in the play. 810 00:48:38,050 --> 00:48:41,675 And he has evidence for all these advantages of play. 811 00:48:45,490 --> 00:48:47,890 And then-- this is interesting-- he 812 00:48:47,890 --> 00:48:50,630 says human research is exploratory behavior 813 00:48:50,630 --> 00:48:53,800 plus play, mostly exploratory behavior. 814 00:48:53,800 --> 00:48:57,010 And human art is mostly playing. 815 00:48:57,010 --> 00:49:00,610 He's not putting it down at all. 816 00:49:00,610 --> 00:49:05,800 He's just pointed out the value of these things in animals. 817 00:49:05,800 --> 00:49:09,370 And now he says, well, if we talk about humans, 818 00:49:09,370 --> 00:49:11,665 this is how he sees human research and art. 819 00:49:17,130 --> 00:49:18,960 Just about all these types of learning, 820 00:49:18,960 --> 00:49:21,190 remember there are species differences, 821 00:49:21,190 --> 00:49:24,770 but we don't know fully all aspects of it. 822 00:49:24,770 --> 00:49:26,510 And what I will talk about next time 823 00:49:26,510 --> 00:49:29,040 is I'll talk a little bit about brain localization. 824 00:49:29,040 --> 00:49:32,180 We can at least make educated guesses in some cases. 825 00:49:32,180 --> 00:49:35,100 Real experiments have been done. 826 00:49:35,100 --> 00:49:39,750 And whether there are any general rules. 827 00:49:39,750 --> 00:49:41,380 The rules seem to be a little different 828 00:49:41,380 --> 00:49:43,720 for all of these different types of learning 829 00:49:43,720 --> 00:49:46,190 that Lorenz has described. 830 00:49:46,190 --> 00:49:53,370 So next time we'll be talking here more about brain pathways. 831 00:49:53,370 --> 00:49:57,440 Neuroscience has made studies, for example, 832 00:49:57,440 --> 00:49:59,670 of hippocampal function and the function 833 00:49:59,670 --> 00:50:04,340 of other parts of the brain-- have given us a different way 834 00:50:04,340 --> 00:50:06,730 to categorize types of learning. 835 00:50:06,730 --> 00:50:09,315 So we'll talk about that next time.