Some useful insights for graduate students beginning their research in ...

Behavioural Brain Research 231 (2012) 234–249

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Review

Some useful insights for graduate students beginning their research in physiological psychology: Anecdotes and attitudes Philip Teitelbaum ∗ Psychology Department, University of Florida, Gainesville, FL 32611, United States

a r t i c l e

i n f o

Article history: Received 10 November 2011 Received in revised form 12 January 2012 Accepted 13 January 2012 Available online 21 January 2012

a b s t r a c t This paper is based on my experiences in 40 years of research in behavioral neuroscience. It is aimed at giving help to beginning graduate students with advice for how to do their research. © 2012 Elsevier B.V. All rights reserved.

Keywords: Physiological psychology Behavioral neuroscience

Contents 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22.

Read old books . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Work with a good scientist . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . My advice on how to pick a problem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . No such thing as failure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Replicate other people’s work . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Psychology is complementary to physiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Simpler to work on the abnormal than on the normal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Don’t rely on statistics to prove that the effect is real . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Use several methods to measure the same thing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stages as a research method . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The value of collaboration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Think with the method of compound opposites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Design a robot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . What is a side-effect? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . How to use the parallel between recovery and development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . To integrate psychology, search for hierarchical levels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . To make an important advance, work on the weakest link . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Use of special language to describe movement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Seek the “elusive obvious” in behavior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Force neurophysiology to confront the complexity of behavior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . To obtain funding, build medicine as well as psychology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Good research can be great fun . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

∗ Tel.: +1 352 392 0615; fax: +1 352 392 7985. E-mail addresses: [email protected], [email protected] 0166-4328/$ – see front matter © 2012 Elsevier B.V. All rights reserved. doi:10.1016/j.bbr.2012.01.030

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“Worship the spirit of criticism. If reduced to itself, it is not an awakener of ideas or a stimulant to great things, but, without it, everything is fallible; it always has the last word. What I am now asking you, and you will ask of your pupils later on, is what is most difficult to an inventor.

P. Teitelbaum / Behavioural Brain Research 231 (2012) 234–249

It is indeed a hard task, when you believe you have found an important scientific fact and are feverishly anxious to publish it, to constrain yourself for days, weeks, years sometimes, to fight with yourself, to try and ruin your own experiments and only to proclaim your discovery after having exhausted all contrary hypotheses.” (Louis Pasteur, quoted in Vallery-Radot, 1923) [1]. This paper is a tribute to the late Eliot Stellar, who helped me get started in research and greatly influenced the rest of my scientific career. It is written as a very personal memoir, describing in a series of anecdotes some advice and attitudes that have helped me shape a personal research style in physiological psychology. Such advice has come from several people, not just from Eliot. However, he was such a constructive, helpful friend and advisor that in his memory I write this, in the hope that the principles in it may be valuable for students beginning their own research in physiological psychology. Some of my attitudes may seem outrageous to some, but I live by them, and explain them to my own students. I hope they will stimulate discussion among students trying to decide how to get their own research off to a good start. 1. Read old books In 1950, shortly after I began my studies as a graduate student in physiological psychology at The John Hopkins University, I became very disheartened. It seemed to me, as a naïve, idealistic and rather arrogant beginner, that my professors in Psychology at Hopkins were not great scientists, and that they could not teach me all I needed to learn. I was mistaken – the professors there at the time, who included Cliff Morgan, Eliot Stellar, Wendell (“Tex”) Garner, Jim Deese, Al Chapanis, Chuck Erickson, Harry Hake, Marv Shaw, and Dick Lazarus were all first-rate.1 In particular, Morgan and Stellar helped me enormously, but at the beginning I did not know that would happen. One night, I was browsing in the library that jointly served the Biology and Psychology Departments, which were then both housed in Mergenthaler Hall on Hopkins Homewood Campus. I had started alphabetically. When I reached “B”, I found an English translation of a book entitled, “An Introduction to the Study of Experimental Medicine” written in 1865 by someone called Claude Bernard [2], of whom I had never heard. I became so excited as I read it that I neglected my studies for days. It cheered me up. I felt that if I could find a few books like that, I might become a good scientist. Only later did I learn that Bernard was a towering genius of scientific research, one of the founders of modern physiology. If one substitutes “psychology” where he says “physiology”, his book is still relevant to physiological psychology [3]. Indeed, Bernard’s method of counter experiment is at the heart of the analysis of stages of recovery that Alan Epstein and I [4] developed as our method of building an understanding of the hierarchy of the organization of behavior. I have never found another book to match his.2

1

The faculty of Hopkins Psychology at that time made sure the incoming students called them by their first names immediately. I was initially uncomfortable with this, but I soon realized that it was a very effective way of getting the students to feel that faculty and students were all equals in the community of scholars working together in the search for new truth. My brother Herman became an undergraduate at Hopkins during the latter half of my graduate studies there. He roomed with me and Pete Lewinsohn, and ate with all the graduate students in Psychology. He was so exhilarated by the atmosphere there that he went on to become a distinguished physiological psychologist, taking a Master’s degree in the joint Physiology-Psychology program at the University of Washington, a Ph.D. in Psychology at McGill University and a postdoctoral fellowship with Roger Sperry at Cal Tech. He currently is retired from the National Institutes of Health. 2 However, some excellent advice for a scientist to live by and a source of great inspiration can be found in “The Life of Pasteur” by Vallery-Radot [1].

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One principle, attributed to Ivan Pavlov, was quoted to me by my mentor at Harvard University, Georg von Bekesy,3 “If you want new ideas, read old books.” Of course, Claude Bernard’s book is an example of the value of this. Reading old books gives you a broader perspective of the field and of the particular problem you are working on. This is difficult to obtain by reading only the latest journals. Therefore, for any subject that you are interested in, read the old books by the great workers. Descartes [5] on the mind-body problem. An understanding of this is critical for how you define your task as a physiological psychologist [3]. Magnus [6] and Sherrington [7] on reflexes, Tinbergen [8] and Lorenz [9] on instinct, Pavlov [10] and Skinner [11] on learning, etc. The problems that they grappled with then are still the central ones of today; the question you must answer for yourself is which of the techniques now available offer any real advance in solving them. 2. Work with a good scientist To develop into a good scientist, work with one. What does such a scientist transmit to his coworkers? Of course, the living example of how he or she works and thinks when faced by each new problem is at the heart of it. However, there is also the store of wisdom that is summed up in personal anecdotes and advice, stimulated by each situation as it arises. My beginning research topic developed from a meeting I had with Cliff Morgan, who was the chairman of the department at the time, and who interviewed and advised all the incoming graduate students. He asked me what I was interested in, and I said, ‘I’m interested in brain and behavior, but I don’t know anything.” He said, ‘Never mind, you just have to jump in.” After some discussion [12], he said, “There are three major unsolved problems in the central nervous system: (1) Recovery of function after damage: How does it occur, since the tissue does not grow back? (2) Release phenomena: How do you get more behavior when you remove part of the nervous system? (3) Delayed appearance of behavior: How come some major effects of damage take as long as six months before they appear? Here are a few references relevant to each of these topics. Go into the library and read about them.” That interview shaped my entire research career. I picked release phenomena as a likely topic to work on. Almost immediately thereafter, I also found a way to study recovery of function in the rat, and have worked on behavioral recovery for the rest of my life. I have always been grateful to Morgan for suggesting those problems. It seems to me, therefore, that one service a mature scientist can do for a beginner is to suggest a problem to work on that will lead to early success in it, and that has enough depth that it can last a lifetime. Following Morgan’s example: let the student start with big issues, and give him or her several choices to work on. 3. My advice on how to pick a problem My general advice to a beginner is: Pick a big, puzzling, abnormal phenomenon in the area you want to work in. Reproduce it in the laboratory or find it in the clinic, and then explore its action under varying conditions that clearly influence its normal counterpart. If you do this, I think there is a good chance that within a short time, you will discover something new. Such phenomena abound in the medical literature about the pathology

3 Bekesy won the Nobel Prize in Medicine for his measurements of the travelling waves on the basilar membrane in the cochlea. Those measurements forced a revision in the thinking about pitch discrimination that had existed since the work of Helmholtz.

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of the nervous system, often labeled with Greek or Latin names, or with the name of the neurologist who first described it. If you love psychology and want to build it, then study the behavioral functions that remain after damage, and how they build on each other during recovery. As explained below, studying recovery will force you to overcome the dominance that medicine has over psychology in the field of physiological psychology, and will force you to build psychology rather than medicine.4 4. No such thing as failure Eliot and I, working side by side, set out to reproduce the release phenomenon5 of hypothalamic hyperphagia in the rat. It had been reported earlier by Hetherington and Ranson [13], and by Brobeck et al. [14]. Eliot and I used his new, personally designed, brass stereotaxis instrument for the rat [15] to produce localized damage in the hypothalamus. Hypothalamic damage was not a method in common use in physiological psychology at that time, so we had to develop our own way of delivering current to damage the tissue, the type of electrode to use, etc. We failed many times. In the course of our repeated failures, Eliot suggested that we vary the locus of our lesions, creating a grid of half millimeter intervals throughout the entire block of tissue that encompassed the medial and lateral hypothalamus as far as the optic chiasm anteriorly and the mammillary bodies posteriorly. In doing so, we reproduced many phenomena reported earlier, including diabetes insipidus, hyperemotionality, and somnolence. Thus, our persistent failures had a silver lining. They enabled us to stumble on the lateral hypothalamic syndrome [16], reported earlier by Anand and Brobeck [17,18]. They also enabled Eliot to speak from first-hand knowledge as he was writing his classic article on “The Physiology of Motivation” [19,20]. This taught me that there really is no such thing as failure when you are doing an experiment. You can usually learn something useful from apparent failure, sometimes more important than the experiment you were trying to do in the first place. In other words, you can turn what seems like a disadvantage into an advantage.

Fig. 1. Hypothalamic obese female rat (right) compared with its normal control. From Teitelbaum [23].

6. Psychology is complementary to physiology

I also learned that it is often very difficult to reproduce a phenomenon reported by another scientist. Many students feel that they must be completely original in their thesis research, and they want to avoid the seemingly dull, unoriginal, task of replication. This is a mistake. Of course, a Ph.D. thesis is much more than mere replication, but this is where you must start if you really want to build your science. Without adequate replication, a field is built on sand. Also, trying to replicate a result reported by others gives you an exquisite appreciation of how seemingly insignificant aspects of method are crucial to reproducing a scientific fact.

I started my research career, under Eliot’s guidance, by studying hyperphagia, in which, after damage in the region of the ventromedial hypothalamic nuclei, rats, people and many other mammals overeat and become enormously obese (see Fig. 1). We soon found that despite their overeating, such animals, particularly after they have become obese, are extremely finicky,6 gobbling up food that tastes good, but rejecting food whose taste and texture is only slightly negative, and which affects hardly at all the intake of the normal animal. Thus, sensory stimuli, such as the taste and texture of the food, exert a great effect on the amount eaten, demonstrating the important role of appetite in the quantitative regulation of food intake [21,22]. [At that time, based on studies of normal animals, it was mistakenly believed by physiologists that taste and appetite affect only the choice of which foods to eat, not the level of their quantitative regulation [23].] This phenomenon also shows that released behavior is always more stimulus-bound, i.e., more reflexive, than normal. As Anand and Brobeck [17,18] had pointed out, after localized bilateral damage in the lateral hypothalamus at the level of the ventromedial nuclei, rats (and many other mammals) stop eating and drinking and will starve to death. They therefore postulated the existence of “hunger centers” just lateral to the inhibitory centers whose destruction produced hyperphagia. However, because Eliot and I were varying the placement of our lesions, some of our lateral hypothalamic lesions damaged the lateral hypothalamus only incompletely, so that many of our animals were aphagic and adipsic for only a few days, and then they recovered by themselves. I was sure that those animals that died would also have recovered if only

4 I use the term “medicine” in the broad sense in which it was used by Claude Bernard [2]. In this usage, it encompasses clinical medicine and all its experimental forms, including physiology, biochemistry, pharmacology, etc. It includes all forms of biological experimentation on living tissue, whether in the context of clinical medicine or not. 5 The term “release phenomenon” is generally taken to mean release from inhibition in the nervous system, centrally, without involving release or imbalance of more peripheral events, as in the gastric or metabolic changes occurring in hypothalamic hyperphagia, which may indeed be playing a major causal role in the overeating. I see no reason for excluding them. It is clear that some form of release is involved, in that the animals overeat after localized damage in the region of the ventromedial nuclei of the hypothalamus. Thus, by removal of the central tissue, an inhibitory influence has been removed, leading to an excess of action involved in eating, whether peripheral or not.

6 When I submitted this paper for publication in the Journal of Comparative and Physiological Psychology (which has since become the Journal of Behavioural Neuroscience), Harry Harlow, the editor, objected to the use of the word “finickiness” on the grounds that it was a colloquialism, and hence not appropriate for a scientific journal. I accepted all his other criticisms, but not that one. I consulted Webster’s Dictionary, and pointed out that Webster classified all words as to whether or not they were colloquial. “Finickiness” was not a colloquialism, as opposed to the word “flabbergast” which was. I went on to say, “Finickiness comes from the word “finical”, meaning to have an exaggeratedly discriminating taste, and therefore it is appropriate to describe not only the behavior of obese hyperphagic rats, but of some consulting editors as well”. Harlow replied, “I can’t quarrel with Webster”, and accepted finickiness. Years later, when I was introduced to him for the first time, he said “Oh! The finickiness man! My wife and I stayed up all night over your reply, but we couldn’t get around Webster.”

5. Replicate other people’s work


we could have kept them alive long enough. We decided to tubefeed all the animals with a liquid diet that Eliot formulated from milk, eggs, a little sugar and some vitamins. He designed it from the viewpoint of physiology (adequate nutrition) rather than psychology (palatability). After some days or weeks after the lesions, the animals looked so normal that I was convinced that they would eat if I could only find a suitably palatable food to offer them. It was the first great thrill of my scientific life when we discovered that such animals, though they refused all else, would eat bits of milk chocolate in sufficient quantity to stay alive without tube-feeding [12,16]. Only years after it became clear from this finding that palatability was vital to the survival of these animals did it occur to me to offer in a dish the liquid diet that I had laboriously tube-fed them three or four times each day, often coming in at 2 a.m. to make sure they got enough to counteract their marked weight loss. Sure enough, that diet was delicious, and, after some recovery, they gobbled it up, regulating their caloric intake on it, despite the fact that they would still totally refuse ordinary rat food and water and die if offered only that.7 This liquid diet gave Dave Williams and me a way of testing whether the animals would eat more of it and regulate their caloric intake on it when water was added to it, and whether they would drink more of it (or even of water alone) in response to dehydration induced by intraperitoneal injection of hypertonic salt solutions [24]. The exaggerated finickiness of hyperphagic and aphagic rats taught me that the psychological view of eating and drinking, in terms of the motivation to eat and the variables that affect appetite, is complementary to the physiological view of homeostatic regulation of caloric intake, leading to very different scientific approaches to the same phenomenon. In other words, each way of looking at the problem is only a partial view – they complete each other. Our psychological way of looking at the problem led Eliot and me to study the motivational and regulatory functions that remained and recovered after the lesions rather than concentrating on the localization of the functions that were missing. It also led us to view lateral hypothalamic electrolytic lesions not in terms of localization, but instead, as partial transections which, because they produce a phenomenon analogous to “spinal shock” [7], temporarily provided all the power of simplification afforded by complete transection, with the added advantage that they permitted us to study the process of resynthesis of function during recovery.8

7 Edward Stricker told me that in the history of the study of salt appetite after adrenalectomy, a similar phenomenon occurred: investigators had known for years that adrenalectomy produced marked salt loss. To counteract this, they tube-fed rats the salt they needed to counteract their salt loss. It was Curt Richter’s insight that such animals would correct their salt deficiency by themselves if they were simply allowed to drink salt solutions voluntarily. 8 Some behavioral neuroscientists consider electrolytic lesions too old-fashioned and unspecific to use in modern neuroscience. This is a perceptually triggered thought-illusion that governs many scientists’ thinking about the techniques being used in an experiment. The illusion is: old = old-fashioned = out of date and inappropriate. Therefore, in order to obtain funds for research, you have to use the most up-to-date (which usually means the most molecular and selective) technique available for manipulating the tissue you wish to remove, despite the fact that this often means you cannot answer adequately the question you are asking, and have to spend a great deal of time and money looking like you are up-to-date, though this may be completely inappropriate for the question at hand. The illusion is reinforced by the idea that you need more and more specificity when you produce a lesion. That is wrong if your objective is not localized but rather partial transection, in order to study the functions that remain and recover. My advice is: Use the most state-ofthe-art technology that will create approval in the peer reviewer; then apply the old, simple methods as well, as you work to find the truth in the phenomenon you wish to understand.

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7. Simpler to work on the abnormal than on the normal Many scientists avoid bizarre abnormalities as phenomena to work on. They say, “First I will understand the normal, then I will work on the abnormal.” This is a thought-illusion. It is based on the instinctive, mistaken inference: bizarre = complex. If the abnormality is produced by loss of function due to damage (i.e., removal) of part of the nervous system, the resultant behavioral phenomena must be simpler than normal, not more complex. The physically simplified remainder of the nervous system cannot produce more complexity than can the whole, intact, system. The reduced system cannot do more with less. Therefore, it is easier to work on the abnormal than on the normal by itself. The abnormal preparation acts like a microscope, in which the action of the fewer remaining variables is magnified, making them easier to study than in the normal. Another thought-illusion combines with bizarre = complex to prevent many scientists from studying remaining function in behavioral abnormalities produced by brain damage. This is the idea that the resulting phenomena are disordered and unlawful [25]. That is not so – the phenomena seen in the abnormal usually have many similarities to those seen at an earlier developmental stage of integration in the normal. They have lost some of the controls that enable the behavior of the normal to be closely regulated and smoothly operating, but what they reveal is quite lawful. My experience is that the remaining variables always act also in the fully intact normal, but their action there is more difficult to isolate. By working on the abnormal, one is simplifying the nervous system by using the method of physical reduction. The power to simplify the remaining behavior function of the nervous system, physically, and to order, has long seemed to me to be the best reason to be a physiological psychologist [3]. The fact that the abnormal is puzzling, to the point of being bizarre, guarantees that any insight you obtain into it will be new and important. By definition, since many scientists avoid the bizarre, working on such a phenomenon eliminates many of the elements of the “rat race” that competition in research can produce, allowing one to work at a leisurely, thoughtful, pace.9 8. Don’t rely on statistics to prove that the effect is real Working on a big phenomenon often eliminates the need to use statistical analysis to decide whether the treatment has produced a reliable effect: this can be obvious in each individual subject. As Smitty Stevens, the great psychophysicist who worked on hearing at Harvard, used to say, “Don’t work down in the noise level.” When I was an instructor at Harvard, I served on Paul Rozin’s advisory committee. He wanted a joint Biology–Psychology degree, so his committee included some biologists. The first thing the psychologists suggested for Rozin to study was statistics. However, a biologist said, “If he learns statistics, he will never do anything good in science.” This echoes the thinking of Claude Bernard [2,26]. My own view on this matter is that if you need inferential statistics to decide whether you have made a difference with your treatment, your method is inadequate. At this point, I have always taken the

9 A friend, at the peak of his scientific career, once said to me, “I can’t take the pace of my work anymore. My knees are giving out.” He had adopted the strategy, accepted by many in physiological psychology, of trying to stay ahead by seizing each new more molecular method that became available, and applying it to his work before it became common in his field. In my opinion, that tactic, though effective, will sooner or later inevitably cause you to feel that you are in a “rat race” in your scientific work. It seems to me to be self-destructive, if not tempered by a strategy that exploits a thought-illusion to eliminate most of the direct competition while carving out your own niche in your field. The latter strategy also gives you time to read old books.

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approach that I must improve my method of measurement or pick a bigger phenomenon to work on. A big phenomenon thus facilitates rapid research. For each experiment, you need to replicate the result in only a few animals or people, rather than having to do so in many to determine statistically whether the variable is having a significant effect. This allows you to vary your method in experiment after experiment, which is far more important in attaining understanding than is the use of many subjects whose results are averaged to produce statistical significance. Personally, I do not depend on statistics to convince me that I have found a real phenomenon. One animal or person is all it should take to do so.10 Sometimes it is difficult to reproduce a phenomenon after I have seen it once. If it is big, however, I know it is real, and that it is important. That is one reason I try always to record each new puzzling, phenomenon on film or video at the time I see it. The video encourages me to keep trying to reproduce it for further analysis. 9. Use several methods to measure the same thing One day at Harvard, I said to von Bekesy, “There seems to me to be a terrible risk in science: you work for years on a problem, publish your results, and then a few years later your results become outdated. How does one guard against that?” He quoted the great chemist, Leibig: “Die Methode ist alles; Alles ist die Methode.” He then said, “I measure everything with five different methods. Only after I have done so do I publish the results. Since most of the variance in any experimental finding is due to the method used to obtain it, I expect my results to stand for about 50 years. By that time, new methods will become available, and new interpretations may result”. I was thunderstruck. Nobody had ever said such a thing to me during my studies in Psychology (though I discovered later that such thinking was common in many other sciences). What was I going to do? I was not a genius like von Bekesy, who could develop five original methods for each experiment. I barely could come up with one method for an experiment, let alone five. After walking around with this for a couple of weeks, I arrived at a compromise: I resolved not to publish any paper until I had measured the phenomenon with two independent methods, despite the pressure on me to publish rapidly.11 Using several methods to measure the same thing is also a good way to strengthen your belief in negative results (when the treatment you have used has no apparent effect). Before you do an experiment, you should always try to build a guarantee against negative results into it. Logically, however, you never really can guarantee them; someone may come along with a better method and show that the treatment really does have an effect, and your negative results go out the window. 10. Stages as a research method In 1957, I began to feel the need to do experiments on cats, not just rats, because I wanted to study the effect of decortications on hypothalamic hyperphagia, and I needed to learn the techniques

10 One person or animal is the appropriate unit for psychology. Therefore, you must be able to see the effects of a given treatment in each animal or person at a time. Studying more than one animal is only necessary to convince yourself and your scientific audience that your idea of what was necessary in the treatment to produce that phenomenon is reliable. 11 The effect was astounding: my reputation in the field grew very rapidly, and many of the papers I published then are still cited today. Best of all, however: my understanding from each experiment deepened, and my work became more satisfying. This is still the best piece of advice I know on how to do good experiments.

of sterile neurosurgery that were necessary in the cat. Complete decortications was relatively easy to accomplish in the cat, but at that time not in the rat [only later did Whishaw and his colleagues [27] perfect this technique for behavioral studies in the rat]. I mentioned my need for a fellowship to Eliot, who merely nodded. However, very soon thereafter, seemingly out of the blue but obviously Eliot’s doing, I received a letter from Frank Beach encouraging me to apply for a Carnegie Foundation Fellowship, which I subsequently received. Eliot had moved to the Neurological Institute at the University of Pennsylvania in 1954, when I took my first job as an instructor at Harvard after my Ph.D. at Johns Hopkins. Along with Jim Sprague, Bill Chambers, and John Liu, Eliot made the conditions so attractive and offered so much friendship and help at the Institute that I decided to spend my fellowship year there in 1958. He also encouraged the late Alan Epstein to spend a postdoctoral fellowship there, after completing his B.A. and M.A. under Eliot’s tutelage on the Hopkins Homewood Campus, and then his M.D. degree at Johns Hopkins Medical School. Alan and I teamed up at the Neurological Institute to study the recovery from lateral hypothalamic aphagia and adipsia. I remember late one evening we were trying to formulate our results for publication, when Alan said something about the stages of recovery in the syndrome. I shouted, “Stages!” The idea of separate stages (see Fig. 2) produced by discrete hierarchically organized transformations during the recovery process transformed our entire understanding of the phenomena we had isolated during the recovery process [4]. I remember vividly when Paul Rozin, Mae Cheng, and I [28] realized that there was a remarkable parallel between stages of recovery and feeding and drinking after lateral hypothalamic damage in the adult rat, as compared to the stages of development in the regulation of food and water intake in the developing normal infant rat (see Fig. 3). This was particularly clear when the rate of development in infancy was slowed down by thyroidectomy at birth [28] or by undernutrition during development [29]. In retrospect, this relationship is obvious: brain damage reduces the hierarchy of the level of integration in the remaining tissue to a lower, more infantile, level. The process of recovery often recapitulates the original levels of integration that were attained during normal development. In the more than 30 years that have passed since its publication, the paper by Epstein and me on the stages of recovery in the lateral hypothalamic syndrome has become one of the top three most cited biologically oriented papers in the hundred year history of the Psychological Review, where it appeared [30]. Stages of recovery offer repeated opportunities to apply Claude Bernard’s powerful method of counterexperiment in the analysis of behavior [3]. They also provide a way of building an understanding of the hierarchy of transformations from simple reflexes to complex operants. One stage = one reflex. 11. The value of collaboration My collaboration with Alan Epstein emphasized for me the tremendous value of the fun, friendship, and increased power that results when you find a congenial person to collaborate with. That person will always have skills and interests that complement your own, so you enhance each other as long as you work together with mutual respect and affection. A good collaboration is like a marriage – it works when you each give wholeheartedly to the partnership. Eliot worked hard to encourage the Psychology Department to offer me a permanent position at Penn, and for the Biology Department to do so for Alan, so we both stayed on there, working together for several years. It is hard to overemphasize the value of collaborating with other scientists and students in your work. You multiply yourself, and produce more, and better, work. I am deeply grateful to the students and colleagues that I have had the opportunity to collaborate with. In that regard, I have found that it pays to be generous


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Fig. 2. Stages of recovery seen in the lateral hypothalamic syndrome. (The critical behavioral events which define the stages are listed on the left.) From Teitelbaum and Epstein [4].

Fig. 3. Comparison of the development of eating and drinking in infancy and their recovery after lateral hypothalamic lesions in the adult. The upper right half of each block represents the recovering lateral hypothalamic rat and the lower-left half the growing thyrodectomized or semi-starved rats (as diagrammed in Fig. 2, except that stage 4, the stage of partial recovery, has been expanded to illustrate the residual defects in eating and drinking). Uniform coloring in each full block indicates similar responses in recovery and development. From Teitelbaum et al. [28].

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with your ideas, your energies, and your resources. What you give comes back a thousand-fold in good will and help from unexpected directions. I have visited research institutes where each laboratory is closed to all its neighbors, and the workers in it are discouraged from talking about their latest ideas, lest someone steal a march on them in the race toward the Nobel Prize. That’s no fun, and it’s not a way to live or work. 12. Think with the method of compound opposites When a physiological psychologist decides to study stages of recovery of function, he or she is placed in the middle of an ambiguity that exists between practicing physiological psychology versus practicing neuroscience. To understand this ambiguity, it is necessary to apply the method of “compound opposites” to your thinking. It is described briefly by Stricker and me [20] in our analysis of the impact on physiological psychology of Eliot Stellar’s article, “The Physiology of Motivation.” In the compound complementarity that exists between structure versus function, and between localization of the tissue that is missing versus behavioral analysis of the functions that remain, concrete tissue structure usually is dominant over considerations of pure abstract function. Here medicine thinks in an opposite way from psychology. Stages of recovery or development are useful if you are trying to build an understanding of the hierarchical organization of behavior by studying the behavioral functions that remain and recover. There is no need to emphasize them when you focus on the tissue functions that are missing and that need to be replaced by a drug or a transplant. Medically oriented neuroscientists who work with brain damage are therefore not typically interested in the behavioral functions that remain and recover, and often cannot understand why you would want to study them. This is the eternal dilemma of the physiological psychologist, who, as Dewsbury [31] points out, always seems to drift into the study of medicine and away from psychology. To avoid this, you must understand the compound opposites that underlie the approach you are taking to the behavioral phenomenon you are trying to understand. There are usually at least four or five intertwined complementarities (opposites) out of a set of twenty or so (they are called themata by Holton [32] who first isolated them) in any given way of thinking about a problem [20]. If, as a physiological psychologist, you are not aware of their existence and don’t understand them, you are likely to be trapped into thinking the way the dominant, usually medically-oriented, people in your field do. 13. Design a robot One way that helps me to guarantee that the principles I derive from my work in physiological psychology are really building psychology, is this: Can my results be applied to the design of a robot as well as to people and animals? If so, I am building an understanding of pure abstract function, which it seems to me is what psychology is all about. This is an old idea: nearly 400 years ago, Descartes derived the concept of the reflex from the example of the automata in Louis IV’s gardens, that rushed out to startle and amuse dalliants who stepped on a concealed treadle as they strolled along. Descartes’ approach is still useful; if you can’t apply the principles and facts that result from your work in physiological psychology to the design of robots, you are probably building medicine rather than psychology [3,20]. A robot is as concrete as an animal or human, even though it has no flesh and blood. Its concreteness helps overcome the complementarity: abstract versus concrete, in which the concreteness of tissue usually reinforces medicine’s dominance over abstract psychological principles in physiological psychology. The state-of-the-art technology involved in robots also matches the technology involved in medicine and makes each approach equally respectable.

Trying to formulate a principle of pure abstract function that can be useful in the design of a robot is an important practical exercise for a physiological psychologist. I wish I had come earlier to this realization. It literally forces you to abstract the general psychological principle that exists in each new behavioral phenomenon, at the time that you discover it. It forces you to rise above the details of the mechanics of the tissue, and to recognize that tissue is not the stuff that mind and behavior are made of. It was only recently, when the Pellises and I were writing a paper describing the stages of reintegration of righting on the ground in lateral hypothalamic rats, that we forced ourselves to come up with the general principle of indirect triggering via allied reflexes to explain the strange fact that at a certain stage of recovery, merely lightly placing the paw on the ground was sufficient to trigger the entire sequence of righting, whereas previously the animal had to push itself over by force, indicating that at the earlier stage the placing reaction, which is a reflex that is allied to righting, was not able by itself to activate the righting sequence. The principle of indirect triggering via allied reflexes explains many phenomena of paradoxical triggering of walking in akinetic people with Parkinson’s disease. It also accounts for many of the phenomena in the stages of recovery of food and water intake in the lateral hypothalamic syndrome [33]. 14. What is a side-effect? A big phenomenon is multi-faceted: it helps you “to see the world in a grain of sand.” For instance, the lateral hypothalamic lesions that produce aphagia and adipsia often also produce catalepsy, akinesia, and other disorders of movement seemingly unrelated to the regulation of eating and drinking, but which can interact with them to decrease intake. They are therefore generally considered to be undesirable “side-effects”, to be avoided by making the lesions as small as possible. However, the thought-illusion that a phenomenon is a side-effect can occur when one takes too narrow a view of what the phenomenon is in the first place. Thus, orienting to stimuli and exploring the environment are part of the act of obtaining food in the animal’s natural habitat. They are deemphasized in the artificially constricted situation of the small cage in which animals are housed when one wishes to study the quantitative aspects of their regulation of food and water intake. Therefore we did the opposite of what everybody else in the field were doing: we deliberately sacrificed the possibility of precise localization of the minimal tissue that must be destroyed to produce the phenomenon. For the sake of a better opportunity for understanding the functions that remained after the damage, we chose to make the lesions larger and larger (see Teitelbaum [12] for our other reasons for doing so). The larger the lesions, the slower the process of recovery, thus permitting us more time to study and experiment with the behavioral phenomena in each stage. This is analogous to Hubbard and Wald’s [34] use of low temperatures to slow down the reversible process of transition from rhodopsin to opsin, thus permitting them to isolate the intermediate stages, in which metaand lumi-rhodopsin are formed. Making larger lesions also was one factor that led us to discover, as shown in Fig. 4, that sensory neglect, a disorder in orientation to sensory stimuli in many sensory modalities, occurs after lateral hypothalamic damage [35,36]. Furthermore, the variables that counteract these “sensorimotor side-effects” also promote eating. For instance, pinching the tail of an akinetic, aphagic, rat can induce it to run away. However, it can also induce an aphagic rat or cat to vigorously eat food placed directly in front of it [36–38]. It will also increase the intake of a normal rat. Therefore, the same variables that control movement also control eating and drinking, and are not really “side-effects” at all [39]. I therefore decided to leave the field of the regulation of food and water intake produced by lateral hypothalamic damage for the


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Fig. 4. A rat with unilateral (right) lateral hypothalamic damage shows precise head orientation and biting to various kinds of stimuli (whisker touch, odor, body touch) on the ipsilateral side (pictures at left) while neglecting the same stimuli presented contralaterally (pictures at right). From Marshall et al. [35].

study of the disorders of movement produced by the same lesions. Because the variables that controlled them were largely the same, it was simpler to observe movement instead of the autonomics of regulation. This is analogous to the exteriorization of autonomic functions that were achieved by Beaumont [40] and by Pavlov [10]. Movement can be observed directly, and by methods that build psychology; autonomics cannot.

15. How to use the parallel between recovery and development With good reason, many scientists instinctively distrust parallels. This is due to the possibility that a parallel is a mere coincidence, without true relevance to the phenomenon under study. One way to guard against accidental coincidence is to study the particular parallel in great detail. As the points of comparison and agreement multiply, it becomes increasingly difficult to dismiss the parallel as mere coincidence. Furthermore, the parallel between recovery and development often has great heuristic value. For instance, David Wolgin and I [39] studied the lateral hypothalamic syndrome in the adult cat. In such animals, we replicated a seemingly strange phenomenon – the bandage-backfall reaction – that had been reported earlier by Van Harreveld and Bogen [41] in otherwise normal cats treated with bulbocapnine, which blocks dopamine function in the brain. In dopaminedeficiency, the independent functional submodules involved in head-scanning, orienting, locomotion, turning, eating, and drinking are blocked [42]. However, the postural support submodule,

which is presumably non-dopaminergic, remains intact and active [43]. The reflexes that comprise the postural support submodule are homeostatic: they defend, maintain and regain upright, stable, immobility. Thus, although it remains immobile when stable and upright, a dopamine-deficient animal will right itself instantly when falling supine in the air, brace against being displaced while standing on the ground [44,45], and cling unmoving in an upright position on a vertical support, such as the back of a chair, or on a pair of vertically separated horizontal bars (see left half of Fig. 5). Seemingly paradoxically, the otherwise immobile dopamine-deficient animal will even leap across the room if it senses instability when its legs begin to slip out from under it [46] (see Fig. 6). Bogen and Van Harreveld showed that if the head and neck are bandaged while the dopamine-deficient cat is clinging upright, the head and neck fall slowly backwards, the forelimbs extend, and the grasp is released, so that the animal falls backward off the chair (see right half of Fig. 5). Wolgin and I replicated this phenomenon in dopamine-deficient cats, rats, and monkeys, and even in a person suffering from Parkinson’s disease (see Fig. 7) [47]. So the phenomenon is quite general, appearing in many species, always based on dopamine-deficiency. Because of the parallel between recovery and development, we immediately tested normal infant kittens, rats, monkey’s, and human babies for the bandage-backfall reaction [48]. Sure enough, the reaction occurs in all of them (see Fig. 8).12 We now believe

12 We submitted these results for publication in Science. One referee proclaimed, “This phenomenon is not within my ken nor that of several colleagues whom I have

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Fig. 5. Left: undrugged adult cat, 2 days after bilateral lateral hypothalamic damage, clinging cataleptically. Right: bandaging the head and neck produces the backfall-reaction. From Teitelbaum et al. [48].

Fig. 6. A jump by a haloperidol-treated rat being pushed forward. At first the rat braces against being pushed forward by shifting its weight backwards (A). When its hind legs begin to slip (B), a leap is triggered (C and D), away from the surface where it is unstable. When it lands on the horizontal table top, it immediately resumes immobility. Thus, cataleptic leaping is merely an allied postural support defensive reflex, triggered by postural instability. From Morrissey et al. [46].

that in the normal adult animal, vestibular input is dominant over tactile input in the control over the position of the head. Dopamine-deficiency reverses this dominance [49]. In the normal animal, when it is clinging to a vertical support, it uses vestibular,

consulted. Furthermore, things that look alike are not necessarily alike. The paper should therefore be rejected.” I called the editor of Science and said, Obviously, the phenomenon is not within the ken of many reviewers – that is why we are submitting it for publication. Furthermore, if two things are alike, how would you expect them to look: similar or different? The editor said, “Hm, yes. I see what you mean.” Nevertheless, the paper was still not accepted by Science. We published it in the Proceedings of the National Academy of Sciences.

kinesthetic, and visual input to sense that its head is being maintained in the correct vertical position. When an elastic bandage is wrapped around its head and neck, the pressure of the bandage supplies a false signal that the head is being supported by a surface. The normal adult animal ignores this signal, and uses the vestibular information to maintain its head upright. In the dopamine-deficient adult, and in the normal infant (in whom dopamine function presumably has not yet developed sufficiently), the dominance relationship is reversed: in the presence of the false signal of support provided by the pressure of the bandage, tactile pressure becomes dominant, and the animal now ignores the vestibular input while it allows the head to sag backwards, and


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Fig. 7. Bandage-backfall reaction in human adult suffering from Parkinson’s disease. From Teitelbaum [47].

releases its grasp, thus falling backward off the supporting surface to which it had been clinging. 16. To integrate psychology, search for hierarchical levels Psychology still suffers from a lack of integration between many of its subfields. One reason for this is that the hierarchical levels of function that are often common to development in infancy and recovery from brain damage in adulthood have not been worked

out yet. Using the parallel between recovery and development as an heuristic technique forces one to work both in developmental and physiological psychology at the same time, thus contributing to the immediate integration of these two subfields [3]. Finally, comparing infants to brain-damaged adults sharpens one’s perception of the details of the behavior of each, more than working on either alone seems to do. That is the main reason that I use this method in my work whenever I can. What is important here is not to prove that the parallel is correct all the time. It often is not. When it is

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Fig. 8. Top left: normal newborn kitten (24 h old) clings to the experimenter’s hand, keeping head and neck erect. Top right: the bandage-backfall reaction in the newborn kitten. Middle left: a two-week old baboon clings to the experimenter’s fingers, keeping head and neck erect. Middle right: bandage-backfall reaction in the infant baboon (from Teitelbaum et al. [48]). Bottom left: 8 week old human infant holds its head and neck erect. Bottom right: bandage-backfall reaction in the same human infant. From Teitelbaum [47].

not, the differences are as illuminating as the similarities. Why the parallel holds sometimes, and why it sometimes does not, seems to me to be an important unanswered question. 17. To make an important advance, work on the weakest link Von Bekesy once mentioned that when he was just starting in his work in the Hungarian equivalent of the Bell Telephone Company, he figured out that the weakest link at that time in telephone

communication was in the degree of reception provided by the telephone receiver. He therefore decided to study the phenomena of hearing, in order to make a contribution to telephone communication. What is the weakest link in the analysis of behavior? In my opinion, the weakest link lies at present in the pure description of behavior. This is due to the fact that many neuroscientists feel that the pure description of behavior is not fundamental, and does not contribute very much to an understanding of how the nervous system works.


One reason for this attitude lies in this complementarity: passive versus active [32]. This ambiguity underlies all experiment. For instance, many physiological psychologists feel the urge to experiment and to intervene actively, by brain damage or stimulation, in the action of the nervous system. To merely observe passively does not seem to them like science. This complementarity exists also in the prejudice against clinical description versus laboratory measurement that has impeded diagnosis in medicine for hundreds of years [25]. However, the main reason for such a negative attitude toward pure description lies in the fact that physiological psychology is different from the rest of psychology because it touches the tissue of the brain. If you don’t touch the tissue of the brain, it is perfectly acceptable merely to observe and describe behavior, as in zoology or psychology in general. Indeed, Lorenz and Tinbergen shared the Nobel prize in Medicine for the pure description of instinctive behavior in animals. As soon as you touch the tissue, however, the ambiguity that is present in physiological psychology crystallizes: if you touch the tissue, then teach me something new about the tissue. That is medicine, not psychology. What is forgotten is that what is being done to the tissue (removal) is for the purpose of simplifying the pure abstract functions that remain. That is psychology, and has nothing to do with tissue. As Claude Bernard [2] said, ‘Mind is to body as time is to a clock.” Time has nothing to do with the hardware being used to measure and reveal it. 18. Use of special language to describe movement Another reason for the deficiency in the pure description of behavior lies in the lack of an adequate language of description of movement in which the basic properties of movement in each part of the body can be defined and analyzed. Levitt and I [50] became aware of the need for such a language when we observed that long after a lateral hypothalamic-damaged rat has recovered the ability to walk around, and seems to explore normally, and nibbles at food in an open field, it can become trapped in a corner, making repetitive head-scanning movements along the floor and walls, but unable to turn around and walk out of the corner. Ilan Golani (an ethologist at Tel Aviv University), David Wolgin, and I [51] applied Eshkol Wachman Movement Notation (EWMN) to this phenomenon, and were able to show that recovery of scanning and exploration after such brain damage occurs at different rates along three continuous dimensions of head-and-body movement (lateral, longitudinal, and vertical). At the stage of recovery of locomotion when forward and lateral snout-scanning movement and forward stepping along the floor had recovered, but upward snoutscanning along the wall of the enclosure had not yet appeared, the rat was trapped in the corner. It could not rear up and turn like a normal rat did, nor could it back out. In order to discover this, we had to recognize the four-fold compound opposite: (1) discrete versus continuous [32]. This involves the choice between description of discrete acts versus continuous dimensions of movement, in which discrete acts are ignored (see Fig. 9). (2) Parts versus whole: We had to combine this with a description of the parts of the animal, rather than the movements of the animal as a whole. (3) Passive versus active: We had to remain passive, observing and filming the animal’s movements for the same period of time each day, making sure not to actively intervene in any way. (4) What is missing versus what is left: we had to focus always on what movements remained and recovered each day, rather than what the animal could not do. 19. Seek the “elusive obvious” in behavior There is another perceptually-triggered thought-illusion that leads to a lack of respect for pure description as science. For

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Fig. 9. Schematic drawings showing a side-view of a rat performing increasingly larger amplitude longitudinal snout scanning movements during three successive phases of recovery. (a) Broken line and full line drawings indicate the extreme positions that the rat assumes during each phase. The cross sign indicates the root of the movement, beyond which there is practically no recruitment of limb and body segments for movement. During increasingly larger longitudinal movements, (b and c) the limb and body segments are recruited in a cephalocaudal order. Drawings were traced from films. From Golani et al. [51].

instance, after applying movement notation to the descriptive analysis of a behavioral phenomenon, you may point out something that has eluded observation for a hundred years. When a cat falls in a supine position in the air, it starts to right itself by turning its head and neck. This is the cervical neck reflex which initiates the act of righting in cats, monkeys, and probably many other animals [6]. To this day, this remains the classic description of righting in the air. However, the rat does not right this way. It rotates its shoulder girdle, carrying the immobile head and neck passively along with it (see Fig. 10). Therefore, air-righting in the rodent is initiated by shoulder rotation, not by the cervical neck reflex [52]. However, once a better description is pointed out, it becomes perceptually so obvious that it is instantly dismissed as trivial, despite the fact that without movement notation, it can remain unobserved for a century in which countless people have looked at it. Perhaps this is what Moshe Feldenkrais [53] meant when he spoke of “the elusive obvious.” There is a way to overcome this perceptually-triggered lack of respect for description. Embody it in a state-of-the-art highly technological method. In science, the latest technology creates automatic respect, reflexively, on the part of peer reviewers. This has some merit, because indeed, science progresses as it creates new methods.13 So in recent years, my coworkers and I have concentrated on developing a computerized method for application of movement notation in the analysis and resynthesis of the description of abnormal gait in Parkinson’s disease. EWMN [54] seems to

13 However, advanced technology is not essential for making an important scientific discovery. When Marshall, Turner, and I discovered sensory neglect in the lateral hypothalamic syndrome, one reviewer commented that the syndrome had became “a whole new ball-game.” I get a lot of pleasure from remembering that we discovered the presence of neglect in aphagia by using a Q-tip cotton swab and a homemade von Frey hair, about a nickel’s worth of technology.

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In the deterioration of parkinsonian gait, there is a regression to an infantile form of weight transfer, in which the weight is shifted forward only after the stepping leg has achieved full contact and support by the ground (see Fig. 11, top). In contrast, the normal adult person shifts the weight forward while there is a groundsupport in only one leg, as indicated by the release of contact of the rear heel from the ground, before the front heel makes contact as the leg steps forward (see Fig. 12) [55]. This is another example of reversal of dominance: as in the bandage-backfall, so too in the gait. Tactile contact (support) becomes dominant over vestibular control, so that both legs have to be in contact with the ground before weight is shifted forward. (From other aspects of the gate, Forssberg et al. [56] had suggested earlier that parkinsonian gait resembled an infantile form of walking). A person with Parkinson’s disease typically walks with short, slow, steps, in a stooped posture, without swinging his arms. However, if we merely ask him to swing his arms when he walks, he instantly automatically walks faster, with longer strides, and with a more erect posture [55,57] (see Fig. 11, bottom). A similar effect occurs if we merely ask him to take larger steps: he swings his arms, walks faster, and uses the more adult form of weight transfer. The principle is: Indirect amplitude modulation via allied reflexes. In other words, voluntarily augmenting the amplitude of one of the allied reflexes involved in walking automatically augments the amplitude of the others. Thus, people with Parkinson’s disease can use these allied reflexes to help themselves to walk better [55,57]. 20. Force neurophysiology to confront the complexity of behavior “For everything there is a reason, and a time for every matter under heaven: . . .. . .. . .. . .. . . a time to break down, and a time to build up; . . .. . .. . .. . .. . ..14

Fig. 10. Tracings from photographs taken from the video monitor showing a righting sequence in which the rat was dropped laterally with respect to the camera. In the first photograph (a), the rat is shown immediately upon release so that its head and feet point skyward. Upon release, the forepaws are tucked up to the sides of the face and then the head, neck, and shoulders rotate en block toward prone (b and c). Once the forequarters are nearly prone, the kindquarters are still 90◦ from prone (d), but then active pelvic rotation brings the remainder of the body to prone (e). This was high-speed video at 1000 frames/s.

The power of Claude Bernard’s method of counterexperiment lies in the fact that he used physical analysis (breaking down) and immediate resynthesis (building up) in each experiment [3]. Unfortunately, science can achieve a great deal of success by continuously oversimplifying without an immediate test for resynthesis. As a physiological psychologist, while studying abnormalities, you can provide a great service to Neuroscience by forcing an awareness that the phenomenon you are studying is a great deal more complicated than is presently “conceived of in their philosophy.” For instance, in the study of the regulation of food and water intake in the 1950s and 1960s, we forced in Physiology an entire reappraisal of the role of taste and appetite, by showing that they were

From Pellis et al. [52].

us to be the best notation system to use, because it is a universal geometrical system for the description of movement, not limited by having been developed for a particular movement style. Also, because EWMN was derived from the analysis of movement in people, it makes sense to study people while trying to computerize the system. So, with some sadness, I have closed my animal laboratory. Now we use movement notation to study the abnormal gait of people afflicted with Parkinson’s disease. As we work toward achieving the computerization of this method, we have found two important facts so far:

14 “For everything there is a reason, and a time for every matter under heaven: a time to be born, a time to die; a time to plant, and a time to pluck up what is planted; a time to kill, and a time to heal; a time to break down, and a time to build up; a time to weep, and a time to laugh; a time to mourn, and a time to dance; a time to cast away stones, and a time to gather stones together; a time to embrace, and a time to refrain from embracing; a time to seek, and a time to lose; a time to keep, and a time to cast away; a time to rend, and a time to sew; a time to keep silence, and a time to speak; a time to love, and a time to hate; a time for war, and a time for peace. From Ecclesiastes, Chapter 3, lines 1–8.


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Fig. 12. A normal adult person shifts the body weight forward while there is ground support in only one leg, as indicated by the release of contact of the rear heel from the ground, before the front heel makes contact as the leg steps forward.

essential for regulation in hypothalamic hyperphagia and vital for survival after lateral hypothalamic damage. Furthermore, by studying recovery rather than concentrating on localization, we showed that lack of drinking was also critical in the syndrome, and that many subtle reflexive, infantile, forms of intake, as in “prandial drinking” were critical for survival after such brain damage [4]. Thus, our work helped to open the field of hypothalamic control of feeding to psychologists. Now they dominate this field. An analogous opportunity for physiological psychologists exists currently in the study of locomotion. In 1981, Cheng and others in our group [58] showed that after damage in the region of the nucleus reticularis tegmenti pontis (NRTP), a remarkable form of galloping locomotion is released in the rat. This form of galloping is greatly dependent on particular sensory stimuli, thus revealing the role of many still unknown reflexes in the control of locomotion. For instance, the NRTP-damaged rat will gallop headlong into a wall. The pressure of the wall on the snout instantly shuts down the galloping, and the rat then sits quietly with its nose in contact with the wall. In an open field, if a rubber band is twisted around the snout of such a rat, simulating the wall by applying localized pressure on the snout, the rat will remain immobile. If the rubber band is snipped, however, the rat with such lesions will instantly gallop forward. Therefore, snout contact clearly inhibits forward locomotion (see also the work of Sinnamon [59] on the role of contact in the inhibition of locomotion).15 Fig. 11. Top: a person with Parkinson’s disease walking naturally. His arms swing very little, and he walks slowly, with relatively short steps. The rear heel does not release contact with the ground until after the front leg has established firm support. So he needs firm support in both legs before he shifts his body weight forward. Bottom: when asked to swing his arms while walking, he immediately walks faster with longer steps, and with head more erect. This improves his shift of weight forward, though in this instance, it is still not fully normal.

15 On a smooth level surface, such a rat will gallop; in a running wheel, however, in which a normal rat will gallop vigorously, the NRTP-damaged rat will only walk slowly and intermittently. Why? We don’t know.

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Haloperidol, which induces akinesia in the normal rat by blocking dopamine receptors in the brain, does not work at all in the NRTP-damaged rat [60]. Nor does morphine. This means that the final common inhibitory pathway by which dopamine-deficiency or morphine inhibits locomotion runs through the area of the NRTP. This also means that a way of reversing the akinesia of Parkinson’s disease can be achieved by developing techniques for inhibiting the action of the NRTP. Drugs that inhibit the action of serotonin, like methysergide, are effective in doing so [61]. An effective use of serotonin inhibition has not yet been worked out as a supplement to l-dopa in the clinical treatment of Parkinson’s disease, although the effects of serotonin blockade in rats recovered from NTRP-damage clearly indicate that such a treatment should help. This phenomenon is crying out for further analysis and exploitation. Maps of the locomotion system make no mention of the inhibitory role of the region of the nucleus reticularis tegmenti pontis [62]. Perhaps this is because these findings do not fit into neurophysiological thinking about locomotion. This offers a great opportunity for brave young physiological psychologists to make an important contribution to this field [see the work of Van Hartesveldt and coworkers [63], and of Sinnamon [59]]. Indeed, the study of such reflexes was largely abandoned after the death of Magnus, perhaps due to the invention of more powerful amplifiers during World War II, so that the young neurophysiologists of that time concentrated on microelectrode work instead. There is a whole field of study of compound, allied, reflexes that is waiting for psychologists to breathe life back into it. 21. To obtain funding, build medicine as well as psychology For a physiological psychologist, building Psychology often means going against the values prevalent among neurologists, and indeed, among many medically-oriented physiological psychologists. Since your research funding depends upon their approval, this is critical to your future success. My advice is to do both – build medicine as well as psychology. Produce important facts about the tissue, as well as important facts about the subcomponents of function. That way you can build Psychology, while continuing to receive approval and financial support from your medicallyoriented peers. However, when you talk about your work, I now think you should not say, “I am working on brain and behavior.” In physiological psychology, it is too easy to dismiss the role of the analysis of behavior with wave of the hand, and instead to concentrate on the details of the mechanics of brain tissue. Say instead, “I am studying the relationship of brain to the hierarchy of behavior.” Hierarchy cannot be waved away; it must be built. 22. Good research can be great fun Although it may look to some as though my work has been all over the map, if you think about it, you will see that it has been the same problem all the time (the one that Cliff Morgan mentioned to me, and that Eliot and Alan started out with me on): how to use recovery of function to discover stages in the lateral hypothalamic syndrome (and syndromes related to it) in order to work out the principles of the hierarchical organization of motivated behavior. Finally, I should point out that perhaps the most important thing that I learned from Eliot Steller is that good research can be great fun. Indeed, I have tried to transmit this to my own students by half-jokingly formulating Teitelbaum’s Law: if there are two methods that can be used for doing an experiment (and there always are), the better method is the one that makes the experiment more fun to do. This may sound frivolous, but it is

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