R

Sp

PP

a

ARRAA

KPB

C

0d

Behavioural Brain Research 231 (2012) 234– 249

Contents lists available at SciVerse ScienceDirect

Behavioural Brain Research

jo u r n al hom epa ge: www.elsev ier .com/ locate /bbr

eview

ome useful insights for graduate students beginning their research inhysiological psychology: Anecdotes and attitudes

hilip Teitelbaum ∗

sychology Department, University of Florida, Gainesville, FL 32611, United States

r t i c l e i n f o

rticle history:eceived 10 November 2011

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 atgiving help to beginning graduate students with advice for how to do their research.

eceived in revised form 12 January 2012ccepted 13 January 2012vailable online 21 January 2012

eywords:hysiological psychology

© 2012 Elsevier B.V. All rights reserved.

ehavioral neuroscience

ontents

1. Read old books . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2352. Work with a good scientist . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2353. My advice on how to pick a problem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2354. No such thing as failure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2365. Replicate other people’s work . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2366. Psychology is complementary to physiology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2367. Simpler to work on the abnormal than on the normal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2378. Don’t rely on statistics to prove that the effect is real . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2379. Use several methods to measure the same thing. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23810. Stages as a research method . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23811. The value of collaboration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23812. Think with the method of compound opposites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24013. Design a robot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24014. What is a side-effect? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24015. How to use the parallel between recovery and development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24116. To integrate psychology, search for hierarchical levels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24317. To make an important advance, work on the weakest link . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24418. Use of special language to describe movement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24519. Seek the “elusive obvious” in behavior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24520. Force neurophysiology to confront the complexity of behavior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24621. To obtain funding, build medicine as well as psychology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248

22. Good research can be great fun . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248

Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

∗ Tel.: +1 352 392 0615; fax: +1 352 392 7985.E-mail addresses: omrainyday@gmail.com, teitelb2@hotmail.com

166-4328/$ – see front matter © 2012 Elsevier B.V. All rights reserved.oi:10.1016/j.bbr.2012.01.030

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248

“Worship the spirit of criticism. If reduced to itself, it is not an

awakener of ideas or a stimulant to great things, but, withoutit, everything is fallible; it always has the last word. What I amnow asking you, and you will ask of your pupils later on, is whatis most difficult to an inventor.

rain R

scoahsIsStuo

1

pvawnwJalw

thhtEBIIggohByooh

dwdwgwoaPcT

i

P. Teitelbaum / Behavioural B

It is indeed a hard task, when you believe you have found animportant scientific fact and are feverishly anxious to publish it,to constrain yourself for days, weeks, years sometimes, to fightwith yourself, to try and ruin your own experiments and onlyto proclaim your discovery after having exhausted all contraryhypotheses.” (Louis Pasteur, quoted in Vallery-Radot, 1923) [1].

This paper is a tribute to the late Eliot Stellar, who helped me gettarted in research and greatly influenced the rest of my scientificareer. It is written as a very personal memoir, describing in a seriesf anecdotes some advice and attitudes that have helped me shape

personal research style in physiological psychology. Such adviceas come from several people, not just from Eliot. However, he wasuch a constructive, helpful friend and advisor that in his memory

write this, in the hope that the principles in it may be valuable fortudents beginning their own research in physiological psychology.ome of my attitudes may seem outrageous to some, but I live byhem, and explain them to my own students. I hope they will stim-late discussion among students trying to decide how to get theirwn research off to a good start.

. Read old books

In 1950, shortly after I began my studies as a graduate student inhysiological psychology at The John Hopkins University, I becameery disheartened. It seemed to me, as a naïve, idealistic and ratherrrogant beginner, that my professors in Psychology at Hopkinsere not great scientists, and that they could not teach me all Ieeded to learn. I was mistaken – the professors there at the time,ho included Cliff Morgan, Eliot Stellar, Wendell (“Tex”) Garner,

im Deese, Al Chapanis, Chuck Erickson, Harry Hake, Marv Shaw,nd Dick Lazarus were all first-rate.1 In particular, Morgan and Stel-ar helped me enormously, but at the beginning I did not know that

ould happen.One night, I was browsing in the library that jointly served

he Biology and Psychology Departments, which were then bothoused in Mergenthaler Hall on Hopkins Homewood Campus. Iad started alphabetically. When I reached “B”, I found an Englishranslation of a book entitled, “An Introduction to the Study ofxperimental Medicine” written in 1865 by someone called Claudeernard [2], of whom I had never heard. I became so excited as

read it that I neglected my studies for days. It cheered me up. felt that if I could find a few books like that, I might become aood scientist. Only later did I learn that Bernard was a toweringenius of scientific research, one of the founders of modern physi-logy. If one substitutes “psychology” where he says “physiology”,is book is still relevant to physiological psychology [3]. Indeed,ernard’s method of counter experiment is at the heart of the anal-

sis of stages of recovery that Alan Epstein and I [4] developed asur method of building an understanding of the hierarchy of therganization of behavior. I have never found another book to matchis.2

1 The faculty of Hopkins Psychology at that time made sure the incoming stu-ents called them by their first names immediately. I was initially uncomfortableith this, but I soon realized that it was a very effective way of getting the stu-ents to feel that faculty and students were all equals in the community of scholarsorking together in the search for new truth. My brother Herman became an under-

raduate at Hopkins during the latter half of my graduate studies there. He roomedith me and Pete Lewinsohn, and ate with all the graduate students in Psychol-

gy. He was so exhilarated by the atmosphere there that he went on to become distinguished physiological psychologist, taking a Master’s degree in the jointhysiology-Psychology program at the University of Washington, a Ph.D. in Psy-hology at McGill University and a postdoctoral fellowship with Roger Sperry at Calech. 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

nspiration can be found in “The Life of Pasteur” by Vallery-Radot [1].

esearch 231 (2012) 234– 249 235

One principle, attributed to Ivan Pavlov, was quoted to me bymy mentor at Harvard University, Georg von Bekesy,3 “If you wantnew ideas, read old books.” Of course, Claude Bernard’s book is anexample of the value of this. Reading old books gives you a broaderperspective of the field and of the particular problem you are work-ing on. This is difficult to obtain by reading only the latest journals.Therefore, for any subject that you are interested in, read the oldbooks by the great workers. Descartes [5] on the mind-body prob-lem. An understanding of this is critical for how you define yourtask as a physiological psychologist [3]. Magnus [6] and Sherring-ton [7] on reflexes, Tinbergen [8] and Lorenz [9] on instinct, Pavlov[10] and Skinner [11] on learning, etc. The problems that they grap-pled with then are still the central ones of today; the question youmust answer for yourself is which of the techniques now availableoffer any real advance in solving them.

2. Work with a good scientist

To develop into a good scientist, work with one. What does sucha scientist transmit to his coworkers? Of course, the living exampleof how he or she works and thinks when faced by each new problemis at the heart of it. However, there is also the store of wisdom thatis summed up in personal anecdotes and advice, stimulated by eachsituation as it arises.

My beginning research topic developed from a meeting I hadwith Cliff Morgan, who was the chairman of the department at thetime, and who interviewed and advised all the incoming graduatestudents. He asked me what I was interested in, and I said, ‘I’minterested in brain and behavior, but I don’t know anything.” Hesaid, ‘Never mind, you just have to jump in.” After some discus-sion [12], he said, “There are three major unsolved problems inthe central nervous system: (1) Recovery of function after dam-age: How does it occur, since the tissue does not grow back? (2)Release phenomena: How do you get more behavior when youremove part of the nervous system? (3) Delayed appearance ofbehavior: How come some major effects of damage take as longas six months before they appear? Here are a few references rel-evant to each of these topics. Go into the library and read aboutthem.”

That interview shaped my entire research career. I pickedrelease phenomena as a likely topic to work on. Almost immedi-ately thereafter, I also found a way to study recovery of functionin the rat, and have worked on behavioral recovery for the rest ofmy life. I have always been grateful to Morgan for suggesting thoseproblems. It seems to me, therefore, that one service a mature sci-entist can do for a beginner is to suggest a problem to work onthat will lead to early success in it, and that has enough depth thatit can last a lifetime. Following Morgan’s example: let the studentstart with big issues, and give him or her several choices to workon.

3. My advice on how to pick a problem

My general advice to a beginner is: Pick a big, puzzling, abnor-mal phenomenon in the area you want to work in. Reproduceit in the laboratory or find it in the clinic, and then explore itsaction under varying conditions that clearly influence its normal

counterpart. If you do this, I think there is a good chance thatwithin a short time, you will discover something new. Suchphenomena abound in the medical literature about the pathology

3 Bekesy won the Nobel Prize in Medicine for his measurements of the travellingwaves on the basilar membrane in the cochlea. Those measurements forced a revi-sion in the thinking about pitch discrimination that had existed since the work ofHelmholtz.

2 rain Research 231 (2012) 234– 249

oolfofpy

4

presicdtootempesmThtfstoa

5

ntwTmwigif

Bfom

tphntoep

36 P. Teitelbaum / Behavioural B

f the nervous system, often labeled with Greek or Latin names,r with the name of the neurologist who first described it. If youove psychology and want to build it, then study the behavioralunctions that remain after damage, and how they build on eachther during recovery. As explained below, studying recovery willorce you to overcome the dominance that medicine has oversychology in the field of physiological psychology, and will forceou to build psychology rather than medicine.4

. No such thing as failure

Eliot and I, working side by side, set out to reproduce the releasehenomenon5 of hypothalamic hyperphagia in the rat. It had beeneported earlier by Hetherington and Ranson [13], and by Brobeckt al. [14]. Eliot and I used his new, personally designed, brasstereotaxis instrument for the rat [15] to produce localized damagen the hypothalamus. Hypothalamic damage was not a method inommon use in physiological psychology at that time, so we had toevelop our own way of delivering current to damage the tissue,he type of electrode to use, etc. We failed many times. In the coursef our repeated failures, Eliot suggested that we vary the locus ofur lesions, creating a grid of half millimeter intervals throughouthe entire block of tissue that encompassed the medial and lat-ral hypothalamus as far as the optic chiasm anteriorly and theammillary bodies posteriorly. In doing so, we reproduced many

henomena reported earlier, including diabetes insipidus, hyper-motionality, and somnolence. Thus, our persistent failures had ailver lining. They enabled us to stumble on the lateral hypothala-ic syndrome [16], reported earlier by Anand and Brobeck [17,18].

hey also enabled Eliot to speak from first-hand knowledge ase was writing his classic article on “The Physiology of Motiva-ion” [19,20]. This taught me that there really is no such thing asailure when you are doing an experiment. You can usually learnomething useful from apparent failure, sometimes more impor-ant than the experiment you were trying to do in the first place. Inther words, you can turn what seems like a disadvantage into andvantage.

. Replicate other people’s work

I also learned that it is often very difficult to reproduce a phe-omenon reported by another scientist. Many students feel thathey must be completely original in their thesis research, and theyant to avoid the seemingly dull, unoriginal, task of replication.

his is a mistake. Of course, a Ph.D. thesis is much more thanere replication, but this is where you must start if you reallyant to build your science. Without adequate replication, a field

s built on sand. Also, trying to replicate a result reported by others

ives you an exquisite appreciation of how seemingly insignif-cant aspects of method are crucial to reproducing a scientificact.

4 I use the term “medicine” in the broad sense in which it was used by Claudeernard [2]. In this usage, it encompasses clinical medicine and all its experimental

orms, including physiology, biochemistry, pharmacology, etc. It includes all formsf biological experimentation on living tissue, whether in the context of clinicaledicine or not.5 The term “release phenomenon” is generally taken to mean release from inhibi-

ion in the nervous system, centrally, without involving release or imbalance of moreeripheral events, as in the gastric or metabolic changes occurring in hypothalamicyperphagia, which may indeed be playing a major causal role in the overeating. I seeo reason for excluding them. It is clear that some form of release is involved, in thathe animals overeat after localized damage in the region of the ventromedial nucleif the hypothalamus. Thus, by removal of the central tissue, an inhibitory influ-nce has been removed, leading to an excess of action involved in eating, whethereripheral or not.

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

6. Psychology is complementary to physiology

I started my research career, under Eliot’s guidance, by studyinghyperphagia, in which, after damage in the region of the ventrome-dial hypothalamic nuclei, rats, people and many other mammalsovereat and become enormously obese (see Fig. 1). We soon foundthat despite their overeating, such animals, particularly after theyhave become obese, are extremely finicky,6 gobbling up food thattastes good, but rejecting food whose taste and texture is onlyslightly negative, and which affects hardly at all the intake of thenormal animal. Thus, sensory stimuli, such as the taste and textureof the food, exert a great effect on the amount eaten, demonstrat-ing the important role of appetite in the quantitative regulation offood intake [21,22]. [At that time, based on studies of normal ani-mals, it was mistakenly believed by physiologists that taste andappetite affect only the choice of which foods to eat, not the levelof their quantitative regulation [23].] This phenomenon also showsthat released behavior is always more stimulus-bound, i.e., morereflexive, than normal.

As Anand and Brobeck [17,18] had pointed out, after localizedbilateral damage in the lateral hypothalamus at the level of the ven-tromedial nuclei, rats (and many other mammals) stop eating anddrinking and will starve to death. They therefore postulated theexistence of “hunger centers” just lateral to the inhibitory centerswhose destruction produced hyperphagia. However, because Eliotand I were varying the placement of our lesions, some of our lat-eral hypothalamic lesions damaged the lateral hypothalamus only

incompletely, so that many of our animals were aphagic and adipsicfor only a few days, and then they recovered by themselves. I wassure that those animals that died would also have recovered if only

6 When I submitted this paper for publication in the Journal of Comparative andPhysiological Psychology (which has since become the Journal of Behavioural Neu-roscience), 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 scientificjournal. I accepted all his other criticisms, but not that one. I consulted Webster’sDictionary, and pointed out that Webster classified all words as to whether or notthey 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 appro-priate to describe not only the behavior of obese hyperphagic rats, but of someconsulting 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! Mywife and I stayed up all night over your reply, but we couldn’t get around Webster.”

rain R

wfmvoaifialOwiogtrstwidi[

tiasaoeEttaib[bmr

attsa

atupuamaciwstow

P. Teitelbaum / Behavioural B

e could have kept them alive long enough. We decided to tube-eed all the animals with a liquid diet that Eliot formulated from

ilk, eggs, a little sugar and some vitamins. He designed it from theiewpoint of physiology (adequate nutrition) rather than psychol-gy (palatability). After some days or weeks after the lesions, thenimals looked so normal that I was convinced that they would eatf I could only find a suitably palatable food to offer them. It was therst great thrill of my scientific life when we discovered that suchnimals, though they refused all else, would eat bits of milk choco-ate in sufficient quantity to stay alive without tube-feeding [12,16].nly years after it became clear from this finding that palatabilityas vital to the survival of these animals did it occur to me to offer

n a dish the liquid diet that I had laboriously tube-fed them threer four times each day, often coming in at 2 a.m. to make sure theyot enough to counteract their marked weight loss. Sure enough,hat diet was delicious, and, after some recovery, they gobbled it up,egulating their caloric intake on it, despite the fact that they wouldtill totally refuse ordinary rat food and water and die if offered onlyhat.7 This liquid diet gave Dave Williams and me a way of testinghether the animals would eat more of it and regulate their caloric

ntake on it when water was added to it, and whether they wouldrink more of it (or even of water alone) in response to dehydration

nduced by intraperitoneal injection of hypertonic salt solutions24].

The exaggerated finickiness of hyperphagic and aphagic ratsaught me that the psychological view of eating and drinking,n terms of the motivation to eat and the variables that affectppetite, is complementary to the physiological view of homeo-tatic regulation of caloric intake, leading to very different scientificpproaches to the same phenomenon. In other words, each wayf looking at the problem is only a partial view – they completeach other. Our psychological way of looking at the problem ledliot and me to study the motivational and regulatory functionshat remained and recovered after the lesions rather than concen-rating on the localization of the functions that were missing. Itlso led us to view lateral hypothalamic electrolytic lesions notn terms of localization, but instead, as partial transections which,ecause they produce a phenomenon analogous to “spinal shock”7], temporarily provided all the power of simplification affordedy complete transection, with the added advantage that they per-itted us to study the process of resynthesis of function during

ecovery.8

7 Edward Stricker told me that in the history of the study of salt appetite afterdrenalectomy, a similar phenomenon occurred: investigators had known for yearshat adrenalectomy produced marked salt loss. To counteract this, they tube-fed ratshe salt they needed to counteract their salt loss. It was Curt Richter’s insight thatuch animals would correct their salt deficiency by themselves if they were simplyllowed to drink salt solutions voluntarily.8 Some behavioral neuroscientists consider electrolytic lesions too old-fashioned

nd unspecific to use in modern neuroscience. This is a perceptually triggeredhought-illusion that governs many scientists’ thinking about the techniques beingsed in an experiment. The illusion is: old = old-fashioned = out of date and inap-ropriate. Therefore, in order to obtain funds for research, you have to use the mostp-to-date (which usually means the most molecular and selective) technique avail-ble for manipulating the tissue you wish to remove, despite the fact that this ofteneans you cannot answer adequately the question you are asking, and have to spend

great deal of time and money looking like you are up-to-date, though this may beompletely inappropriate for the question at hand. The illusion is reinforced by thedea that you need more and more specificity when you produce a lesion. That is

rong if your objective is not localized but rather partial transection, in order totudy the functions that remain and recover. My advice is: Use the most state-of-he-art technology that will create approval in the peer reviewer; then apply theld, simple methods as well, as you work to find the truth in the phenomenon youish to understand.

esearch 231 (2012) 234– 249 237

7. Simpler to work on the abnormal than on the normal

Many scientists avoid bizarre abnormalities as phenomena towork on. They say, “First I will understand the normal, then I willwork on the abnormal.” This is a thought-illusion. It is based on theinstinctive, mistaken inference: bizarre = complex. If the abnormal-ity is produced by loss of function due to damage (i.e., removal) ofpart of the nervous system, the resultant behavioral phenomenamust be simpler than normal, not more complex. The physicallysimplified remainder of the nervous system cannot produce morecomplexity than can the whole, intact, system. The reduced sys-tem cannot do more with less. Therefore, it is easier to work on theabnormal than on the normal by itself. The abnormal preparationacts like a microscope, in which the action of the fewer remainingvariables is magnified, making them easier to study than in thenormal.

Another thought-illusion combines with bizarre = complex toprevent many scientists from studying remaining function inbehavioral abnormalities produced by brain damage. This is theidea that the resulting phenomena are disordered and unlawful[25]. That is not so – the phenomena seen in the abnormal usuallyhave many similarities to those seen at an earlier developmentalstage of integration in the normal. They have lost some of the con-trols that enable the behavior of the normal to be closely regulatedand smoothly operating, but what they reveal is quite lawful.

My experience is that the remaining variables always act alsoin the fully intact normal, but their action there is more difficultto isolate. By working on the abnormal, one is simplifying the ner-vous system by using the method of physical reduction. The powerto simplify the remaining behavior function of the nervous system,physically, and to order, has long seemed to me to be the best reasonto be a physiological psychologist [3]. The fact that the abnormal ispuzzling, to the point of being bizarre, guarantees that any insightyou obtain into it will be new and important. By definition, sincemany scientists avoid the bizarre, working on such a phenomenoneliminates many of the elements of the “rat race” that competi-tion 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 usestatistical analysis to decide whether the treatment has produceda reliable effect: this can be obvious in each individual subject. AsSmitty Stevens, the great psychophysicist who worked on hearingat Harvard, used to say, “Don’t work down in the noise level.”

When I was an instructor at Harvard, I served on Paul Rozin’sadvisory committee. He wanted a joint Biology–Psychology degree,so his committee included some biologists. The first thing the psy-chologists suggested for Rozin to study was statistics. However, abiologist said, “If he learns statistics, he will never do anything goodin science.” This echoes the thinking of Claude Bernard [2,26]. My

own view on this matter is that if you need inferential statistics todecide 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 thepace 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 seizingeach new more molecular method that became available, and applying it to his workbefore 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 yourscientific work. It seems to me to be self-destructive, if not tempered by a strategythat exploits a thought-illusion to eliminate most of the direct competition whilecarving out your own niche in your field. The latter strategy also gives you time toread old books.

2 rain R

aa

eocyios

htatnTa

9

bloctaaotm

dsgfiwaInm

wmengan

1

jh

mtyp

mms

38 P. Teitelbaum / Behavioural B

pproach that I must improve my method of measurement or pick bigger phenomenon to work on.

A big phenomenon thus facilitates rapid research. For eachxperiment, you need to replicate the result in only a few animalsr people, rather than having to do so in many to determine statisti-ally whether the variable is having a significant effect. This allowsou to vary your method in experiment after experiment, whichs far more important in attaining understanding than is the usef many subjects whose results are averaged to produce statisticalignificance.

Personally, I do not depend on statistics to convince me that Iave found a real phenomenon. One animal or person is all it shouldake to do so.10 Sometimes it is difficult to reproduce a phenomenonfter I have seen it once. If it is big, however, I know it is real, andhat it is important. That is one reason I try always to record eachew puzzling, phenomenon on film or video at the time I see it.he video encourages me to keep trying to reproduce it for furthernalysis.

. Use several methods to measure the same thing

One day at Harvard, I said to von Bekesy, “There seems to me toe a terrible risk in science: you work for years on a problem, pub-

ish your results, and then a few years later your results becomeutdated. How does one guard against that?” He quoted the greathemist, Leibig: “Die Methode ist alles; Alles ist die Methode.” Hehen said, “I measure everything with five different methods. Onlyfter I have done so do I publish the results. Since most of the vari-nce in any experimental finding is due to the method used tobtain it, I expect my results to stand for about 50 years. By thatime, new methods will become available, and new interpretations

ay result”.I was thunderstruck. Nobody had ever said such a thing to me

uring my studies in Psychology (though I discovered later thatuch thinking was common in many other sciences). What was Ioing to do? I was not a genius like von Bekesy, who could developve original methods for each experiment. I barely could come upith one method for an experiment, let alone five. After walking

round with this for a couple of weeks, I arrived at a compromise: resolved not to publish any paper until I had measured the phe-omenon with two independent methods, despite the pressure one to publish rapidly.11

Using several methods to measure the same thing is also a gooday to strengthen your belief in negative results (when the treat-ent you have used has no apparent effect). Before you do an

xperiment, you should always try to build a guarantee againstegative results into it. Logically, however, you never really canuarantee them; someone may come along with a better methodnd show that the treatment really does have an effect, and youregative results go out the window.

0. Stages as a research method

In 1957, I began to feel the need to do experiments on cats, notust rats, because I wanted to study the effect of decortications onypothalamic hyperphagia, and I needed to learn the techniques

10 One person or animal is the appropriate unit for psychology. Therefore, youust be able to see the effects of a given treatment in each animal or person at a

ime. Studying more than one animal is only necessary to convince yourself andour scientific audience that your idea of what was necessary in the treatment toroduce that phenomenon is reliable.11 The effect was astounding: my reputation in the field grew very rapidly, andany of the papers I published then are still cited today. Best of all, however:y understanding from each experiment deepened, and my work became more

atisfying. This is still the best piece of advice I know on how to do good experiments.

esearch 231 (2012) 234– 249

of sterile neurosurgery that were necessary in the cat. Completedecortications was relatively easy to accomplish in the cat, butat that time not in the rat [only later did Whishaw and his col-leagues [27] perfect this technique for behavioral studies in therat]. I mentioned my need for a fellowship to Eliot, who merelynodded. However, very soon thereafter, seemingly out of the bluebut obviously Eliot’s doing, I received a letter from Frank Beachencouraging me to apply for a Carnegie Foundation Fellowship,which I subsequently received. Eliot had moved to the NeurologicalInstitute at the University of Pennsylvania in 1954, when I took myfirst job as an instructor at Harvard after my Ph.D. at Johns Hopkins.Along with Jim Sprague, Bill Chambers, and John Liu, Eliot made theconditions so attractive and offered so much friendship and help atthe Institute that I decided to spend my fellowship year there in1958. He also encouraged the late Alan Epstein to spend a post-doctoral fellowship there, after completing his B.A. and M.A. underEliot’s tutelage on the Hopkins Homewood Campus, and then hisM.D. degree at Johns Hopkins Medical School. Alan and I teamedup at the Neurological Institute to study the recovery from lateralhypothalamic aphagia and adipsia. I remember late one evening wewere trying to formulate our results for publication, when Alan saidsomething about the stages of recovery in the syndrome. I shouted,“Stages!” The idea of separate stages (see Fig. 2) produced by dis-crete hierarchically organized transformations during the recoveryprocess transformed our entire understanding of the phenomenawe 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 ofrecovery and feeding and drinking after lateral hypothalamic dam-age in the adult rat, as compared to the stages of development in theregulation of food and water intake in the developing normal infantrat (see Fig. 3). This was particularly clear when the rate of develop-ment in infancy was slowed down by thyroidectomy at birth [28] orby undernutrition during development [29]. In retrospect, this rela-tionship is obvious: brain damage reduces the hierarchy of the levelof integration in the remaining tissue to a lower, more infantile,level. The process of recovery often recapitulates the original levelsof 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 lat-eral hypothalamic syndrome has become one of the top three mostcited biologically oriented papers in the hundred year history ofthe Psychological Review, where it appeared [30]. Stages of recov-ery offer repeated opportunities to apply Claude Bernard’s powerfulmethod of counterexperiment in the analysis of behavior [3]. Theyalso provide a way of building an understanding of the hierarchyof transformations from simple reflexes to complex operants. Onestage = one reflex.

11. The value of collaboration

My collaboration with Alan Epstein emphasized for me thetremendous value of the fun, friendship, and increased power thatresults when you find a congenial person to collaborate with. Thatperson will always have skills and interests that complement yourown, so you enhance each other as long as you work together withmutual 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 offerme a permanent position at Penn, and for the Biology Departmentto do so for Alan, so we both stayed on there, working together forseveral years. It is hard to overemphasize the value of collaborating

with other scientists and students in your work. You multiply your-self, and produce more, and better, work. I am deeply grateful to thestudents and colleagues that I have had the opportunity to collab-orate with. In that regard, I have found that it pays to be generous

P. Teitelbaum / Behavioural Brain Research 231 (2012) 234– 249 239

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 blockrepresents 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 inrecovery and development.

From Teitelbaum et al. [28].

2 rain R

wcdifta

1

ripsIiPetfoimhfttenrtgtyatt(ilut

1

fcaspDtaapcrhotoTtr

40 P. Teitelbaum / Behavioural B

ith your ideas, your energies, and your resources. What you giveomes back a thousand-fold in good will and help from unexpectedirections. I have visited research institutes where each laboratory

s closed to all its neighbors, and the workers in it are discouragedrom talking about their latest ideas, lest someone steal a march onhem in the race toward the Nobel Prize. That’s no fun, and it’s not

way to live or work.

2. Think with the method of compound opposites

When a physiological psychologist decides to study stages ofecovery of function, he or she is placed in the middle of an ambigu-ty that exists between practicing physiological psychology versusracticing neuroscience. To understand this ambiguity, it is neces-ary to apply the method of “compound opposites” to your thinking.t is described briefly by Stricker and me [20] in our analysis of thempact on physiological psychology of Eliot Stellar’s article, “Thehysiology of Motivation.” In the compound complementarity thatxists between structure versus function, and between localiza-ion of the tissue that is missing versus behavioral analysis of theunctions that remain, concrete tissue structure usually is dominantver considerations of pure abstract function. Here medicine thinksn an opposite way from psychology. Stages of recovery or develop-

ent are useful if you are trying to build an understanding of theierarchical organization of behavior by studying the behavioral

unctions that remain and recover. There is no need to emphasizehem when you focus on the tissue functions that are missing andhat need to be replaced by a drug or a transplant. Medically ori-nted neuroscientists who work with brain damage are thereforeot typically interested in the behavioral functions that remain andecover, and often cannot understand why you would want to studyhem. This is the eternal dilemma of the physiological psycholo-ist, who, as Dewsbury [31] points out, always seems to drift intohe study of medicine and away from psychology. To avoid this,ou must understand the compound opposites that underlie thepproach you are taking to the behavioral phenomenon you arerying to understand. There are usually at least four or five inter-wined complementarities (opposites) out of a set of twenty or sothey are called themata by Holton [32] who first isolated them)n any given way of thinking about a problem [20]. If, as a physio-ogical psychologist, you are not aware of their existence and don’tnderstand them, you are likely to be trapped into thinking the wayhe dominant, usually medically-oriented, people in your field do.

3. Design a robot

One way that helps me to guarantee that the principles I deriverom my work in physiological psychology are really building psy-hology, is this: Can my results be applied to the design of a robots well as to people and animals? If so, I am building an under-tanding of pure abstract function, which it seems to me is whatsychology is all about. This is an old idea: nearly 400 years ago,escartes derived the concept of the reflex from the example of

he automata in Louis IV’s gardens, that rushed out to startle andmuse dalliants who stepped on a concealed treadle as they strolledlong. Descartes’ approach is still useful; if you can’t apply therinciples and facts that result from your work in physiological psy-hology to the design of robots, you are probably building medicineather than psychology [3,20]. A robot is as concrete as an animal oruman, even though it has no flesh and blood. Its concreteness helpsvercome the complementarity: abstract versus concrete, in whichhe concreteness of tissue usually reinforces medicine’s dominance

ver abstract psychological principles in physiological psychology.he state-of-the-art technology involved in robots also matches theechnology involved in medicine and makes each approach equallyespectable.

esearch 231 (2012) 234– 249

Trying to formulate a principle of pure abstract function that canbe useful in the design of a robot is an important practical exercisefor a physiological psychologist. I wish I had come earlier to thisrealization. It literally forces you to abstract the general psycholog-ical principle that exists in each new behavioral phenomenon, atthe time that you discover it. It forces you to rise above the detailsof the mechanics of the tissue, and to recognize that tissue is notthe stuff that mind and behavior are made of. It was only recently,when the Pellises and I were writing a paper describing the stagesof reintegration of righting on the ground in lateral hypothalamicrats, that we forced ourselves to come up with the general principleof indirect triggering via allied reflexes to explain the strange factthat at a certain stage of recovery, merely lightly placing the paw onthe ground was sufficient to trigger the entire sequence of righting,whereas previously the animal had to push itself over by force, indi-cating that at the earlier stage the placing reaction, which is a reflexthat is allied to righting, was not able by itself to activate the right-ing sequence. The principle of indirect triggering via allied reflexesexplains many phenomena of paradoxical triggering of walking inakinetic people with Parkinson’s disease. It also accounts for manyof the phenomena in the stages of recovery of food and water intakein the lateral hypothalamic syndrome [33].

14. What is a side-effect?

A big phenomenon is multi-faceted: it helps you “to see theworld in a grain of sand.” For instance, the lateral hypothala-mic lesions that produce aphagia and adipsia often also producecatalepsy, akinesia, and other disorders of movement seeminglyunrelated to the regulation of eating and drinking, but which caninteract with them to decrease intake. They are therefore generallyconsidered to be undesirable “side-effects”, to be avoided by mak-ing the lesions as small as possible. However, the thought-illusionthat a phenomenon is a side-effect can occur when one takes toonarrow a view of what the phenomenon is in the first place. Thus,orienting to stimuli and exploring the environment are part of theact of obtaining food in the animal’s natural habitat. They are deem-phasized in the artificially constricted situation of the small cage inwhich animals are housed when one wishes to study the quantita-tive aspects of their regulation of food and water intake. Thereforewe did the opposite of what everybody else in the field were doing:we deliberately sacrificed the possibility of precise localization ofthe minimal tissue that must be destroyed to produce the phe-nomenon. For the sake of a better opportunity for understandingthe functions that remained after the damage, we chose to makethe lesions larger and larger (see Teitelbaum [12] for our other rea-sons for doing so). The larger the lesions, the slower the process ofrecovery, thus permitting us more time to study and experimentwith the behavioral phenomena in each stage. This is analogous toHubbard and Wald’s [34] use of low temperatures to slow down thereversible process of transition from rhodopsin to opsin, thus per-mitting them to isolate the intermediate stages, in which meta-and lumi-rhodopsin are formed. Making larger lesions also wasone factor that led us to discover, as shown in Fig. 4, that sensoryneglect, a disorder in orientation to sensory stimuli in many sensorymodalities, occurs after lateral hypothalamic damage [35,36].

Furthermore, the variables that counteract these “sensorimotorside-effects” also promote eating. For instance, pinching the tailof an akinetic, aphagic, rat can induce it to run away. However, itcan also induce an aphagic rat or cat to vigorously eat food placeddirectly in front of it [36–38]. It will also increase the intake of anormal rat. Therefore, the same variables that control movement

also control eating and drinking, and are not really “side-effects” atall [39].

I therefore decided to leave the field of the regulation of foodand water intake produced by lateral hypothalamic damage for the

P. Teitelbaum / Behavioural Brain Research 231 (2012) 234– 249 241

F d oriet ntrala

F

sBirfMp

1d

acstsdlvhcrBwdhi

we immediately tested normal infant kittens, rats, monkey’s, andhuman babies for the bandage-backfall reaction [48]. Sure enough,the reaction occurs in all of them (see Fig. 8).12 We now believe

ig. 4. A rat with unilateral (right) lateral hypothalamic damage shows precise heahe ipsilateral side (pictures at left) while neglecting the same stimuli presented co

rom Marshall et al. [35].

tudy of the disorders of movement produced by the same lesions.ecause the variables that controlled them were largely the same,

t was simpler to observe movement instead of the autonomics ofegulation. This is analogous to the exteriorization of autonomicunctions that were achieved by Beaumont [40] and by Pavlov [10].

ovement can be observed directly, and by methods that buildsychology; autonomics cannot.

5. How to use the parallel between recovery andevelopment

With good reason, many scientists instinctively distrust par-llels. This is due to the possibility that a parallel is a mereoincidence, without true relevance to the phenomenon undertudy. One way to guard against accidental coincidence is to studyhe particular parallel in great detail. As the points of compari-on and agreement multiply, it becomes increasingly difficult toismiss the parallel as mere coincidence. Furthermore, the paral-

el between recovery and development often has great heuristicalue. For instance, David Wolgin and I [39] studied the lateralypothalamic syndrome in the adult cat. In such animals, we repli-ated a seemingly strange phenomenon – the bandage-backfalleaction – that had been reported earlier by Van Harreveld andogen [41] in otherwise normal cats treated with bulbocapnine,

hich blocks dopamine function in the brain. In dopamine-eficiency, the independent functional submodules involved inead-scanning, orienting, locomotion, turning, eating, and drink-

ng are blocked [42]. However, the postural support submodule,

ntation and biting to various kinds of stimuli (whisker touch, odor, body touch) onterally (pictures at right).

which is presumably non-dopaminergic, remains intact and active[43]. The reflexes that comprise the postural support submoduleare homeostatic: they defend, maintain and regain upright, sta-ble, immobility. Thus, although it remains immobile when stableand upright, a dopamine-deficient animal will right itself instantlywhen falling supine in the air, brace against being displaced whilestanding on the ground [44,45], and cling unmoving in an uprightposition on a vertical support, such as the back of a chair, or on a pairof vertically separated horizontal bars (see left half of Fig. 5). Seem-ingly paradoxically, the otherwise immobile dopamine-deficientanimal will even leap across the room if it senses instability whenits legs begin to slip out from under it [46] (see Fig. 6). Bogen andVan Harreveld showed that if the head and neck are bandagedwhile the dopamine-deficient cat is clinging upright, the head andneck fall slowly backwards, the forelimbs extend, and the graspis released, so that the animal falls backward off the chair (seeright half of Fig. 5). Wolgin and I replicated this phenomenon indopamine-deficient cats, rats, and monkeys, and even in a personsuffering from Parkinson’s disease (see Fig. 7) [47]. So the phe-nomenon is quite general, appearing in many species, always basedon dopamine-deficiency.

Because of the parallel between recovery and development,

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

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

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 hindl unstaT red by

F

toDa

cstsemi

egs begin to slip (B), a leap is triggered (C and D), away from the surface where it ishus, cataleptic leaping is merely an allied postural support defensive reflex, trigge

rom Morrissey et al. [46].

hat in the normal adult animal, vestibular input is dominant

ver tactile input in the control over the position of the head.opamine-deficiency reverses this dominance [49]. In the normalnimal, when it is clinging to a vertical support, it uses vestibular,

onsulted. Furthermore, things that look alike are not necessarily alike. The paperhould therefore be rejected.” I called the editor of Science and said, Obviously,he phenomenon is not within the ken of many reviewers – that is why we areubmitting it for publication. Furthermore, if two things are alike, how would youxpect them to look: similar or different? The editor said, “Hm, yes. I see what youean.” Nevertheless, the paper was still not accepted by Science. We published it

n the Proceedings of the National Academy of Sciences.

ble. When it lands on the horizontal table top, it immediately resumes immobility. postural instability.

kinesthetic, and visual input to sense that its head is being main-tained in the correct vertical position. When an elastic bandageis wrapped around its head and neck, the pressure of the ban-dage supplies a false signal that the head is being supported bya surface. The normal adult animal ignores this signal, and usesthe vestibular information to maintain its head upright. In thedopamine-deficient adult, and in the normal infant (in whomdopamine function presumably has not yet developed sufficiently),

the dominance relationship is reversed: in the presence of the falsesignal of support provided by the pressure of the bandage, tac-tile pressure becomes dominant, and the animal now ignores thevestibular input while it allows the head to sag backwards, and

P. Teitelbaum / Behavioural Brain Research 231 (2012) 234– 249 243

an aF

rt

1

oor

Fig. 7. Bandage-backfall reaction in humrom Teitelbaum [47].

eleases its grasp, thus falling backward off the supporting surfaceo which it had been clinging.

6. To integrate psychology, search for hierarchical levels

Psychology still suffers from a lack of integration between manyf its subfields. One reason for this is that the hierarchical levelsf function that are often common to development in infancy andecovery from brain damage in adulthood have not been worked

dult suffering from Parkinson’s disease.

out yet. Using the parallel between recovery and development asan heuristic technique forces one to work both in developmentaland physiological psychology at the same time, thus contributing tothe immediate integration of these two subfields [3]. Finally, com-paring infants to brain-damaged adults sharpens one’s perception

of the details of the behavior of each, more than working on eitheralone seems to do. That is the main reason that I use this methodin my work whenever I can. What is important here is not to provethat the parallel is correct all the time. It often is not. When it is

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

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 newbornkitten. 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( nd ne

F

npt

1l

hp

from Teitelbaum et al. [48]). Bottom left: 8 week old human infant holds its head a

rom Teitelbaum [47].

ot, the differences are as illuminating as the similarities. Why thearallel holds sometimes, and why it sometimes does not, seemso me to be an important unanswered question.

7. To make an important advance, work on the weakestink

Von Bekesy once mentioned that when he was just starting inis work in the Hungarian equivalent of the Bell Telephone Com-any, he figured out that the weakest link at that time in telephone

ck erect. Bottom right: bandage-backfall reaction in the same human infant.

communication was in the degree of reception provided by the tele-phone receiver. He therefore decided to study the phenomena ofhearing, in order to make a contribution to telephone communica-tion.

What is the weakest link in the analysis of behavior? In myopinion, the weakest link lies at present in the pure description

of behavior. This is due to the fact that many neuroscientists feelthat the pure description of behavior is not fundamental, and doesnot contribute very much to an understanding of how the nervoussystem works.

rain Research 231 (2012) 234– 249 245

simtntsy

pftaotbaitwpaih

1

bopbttbmwGItaa(mfsrawvtwWrvaswa

1

l

Fig. 9. Schematic drawings showing a side-view of a rat performing increasinglylarger amplitude longitudinal snout scanning movements during three successivephases of recovery. (a) Broken line and full line drawings indicate the extreme posi-tions that the rat assumes during each phase. The cross sign indicates the root of themovement, beyond which there is practically no recruitment of limb and body seg-ments for movement. During increasingly larger longitudinal movements, (b and c)

centrated on developing a computerized method for application ofmovement notation in the analysis and resynthesis of the descrip-tion of abnormal gait in Parkinson’s disease. EWMN [54] seems to

13 However, advanced technology is not essential for making an important sci-

P. Teitelbaum / Behavioural B

One reason for this attitude lies in this complementarity: pas-ive versus active [32]. This ambiguity underlies all experiment. Fornstance, many physiological psychologists feel the urge to experi-

ent and to intervene actively, by brain damage or stimulation, inhe action of the nervous system. To merely observe passively doesot seem to them like science. This complementarity exists also inhe prejudice against clinical description versus laboratory mea-urement that has impeded diagnosis in medicine for hundreds ofears [25].

However, the main reason for such a negative attitude towardure description lies in the fact that physiological psychology is dif-erent from the rest of psychology because it touches the tissue ofhe brain. If you don’t touch the tissue of the brain, it is perfectlycceptable merely to observe and describe behavior, as in zool-gy or psychology in general. Indeed, Lorenz and Tinbergen sharedhe Nobel prize in Medicine for the pure description of instinctiveehavior in animals. As soon as you touch the tissue, however, thembiguity that is present in physiological psychology crystallizes:f you touch the tissue, then teach me something new about theissue. That is medicine, not psychology. What is forgotten is thathat is being done to the tissue (removal) is for the purpose of sim-lifying the pure abstract functions that remain. That is psychology,nd has nothing to do with tissue. As Claude Bernard [2] said, ‘Minds to body as time is to a clock.” Time has nothing to do with theardware being used to measure and reveal it.

8. Use of special language to describe movement

Another reason for the deficiency in the pure description ofehavior lies in the lack of an adequate language of descriptionf movement in which the basic properties of movement in eachart of the body can be defined and analyzed. Levitt and I [50]ecame aware of the need for such a language when we observedhat long after a lateral hypothalamic-damaged rat has recoveredhe ability to walk around, and seems to explore normally, and nib-les at food in an open field, it can become trapped in a corner,aking repetitive head-scanning movements along the floor andalls, but unable to turn around and walk out of the corner. Ilanolani (an ethologist at Tel Aviv University), David Wolgin, and

[51] applied Eshkol Wachman Movement Notation (EWMN) tohis phenomenon, and were able to show that recovery of scanningnd exploration after such brain damage occurs at different rateslong three continuous dimensions of head-and-body movementlateral, longitudinal, and vertical). At the stage of recovery of loco-

otion when forward and lateral snout-scanning movement andorward stepping along the floor had recovered, but upward snout-canning along the wall of the enclosure had not yet appeared, theat was trapped in the corner. It could not rear up and turn like

normal rat did, nor could it back out. In order to discover this,e had to recognize the four-fold compound opposite: (1) discrete

ersus continuous [32]. This involves the choice between descrip-ion of discrete acts versus continuous dimensions of movement, inhich discrete acts are ignored (see Fig. 9). (2) Parts versus whole:e had to combine this with a description of the parts of the animal,

ather than the movements of the animal as a whole. (3) Passiveersus active: We had to remain passive, observing and filming thenimal’s movements for the same period of time each day, makingure not to actively intervene in any way. (4) What is missing versushat is left: we had to focus always on what movements remained

nd recovered each day, rather than what the animal could not do.

9. Seek the “elusive obvious” in behavior

There is another perceptually-triggered thought-illusion thateads to a lack of respect for pure description as science. For

the limb and body segments are recruited in a cephalocaudal order. Drawings weretraced from films.

From Golani et al. [51].

instance, after applying movement notation to the descriptive anal-ysis of a behavioral phenomenon, you may point out something thathas eluded observation for a hundred years. When a cat falls in asupine position in the air, it starts to right itself by turning its headand neck. This is the cervical neck reflex which initiates the act ofrighting in cats, monkeys, and probably many other animals [6].To this day, this remains the classic description of righting in theair. However, the rat does not right this way. It rotates its shouldergirdle, carrying the immobile head and neck passively along withit (see Fig. 10). Therefore, air-righting in the rodent is initiated byshoulder rotation, not by the cervical neck reflex [52]. However,once a better description is pointed out, it becomes perceptuallyso obvious that it is instantly dismissed as trivial, despite the factthat without movement notation, it can remain unobserved for acentury in which countless people have looked at it. Perhaps this iswhat Moshe Feldenkrais [53] meant when he spoke of “the elusiveobvious.”

There is a way to overcome this perceptually-triggered lackof respect for description. Embody it in a state-of-the-art highlytechnological method. In science, the latest technology createsautomatic respect, reflexively, on the part of peer reviewers. Thishas some merit, because indeed, science progresses as it createsnew methods.13 So in recent years, my coworkers and I have con-

entific discovery. When Marshall, Turner, and I discovered sensory neglect in thelateral hypothalamic syndrome, one reviewer commented that the syndrome hadbecame “a whole new ball-game.” I get a lot of pleasure from remembering that wediscovered the presence of neglect in aphagia by using a Q-tip cotton swab and ahomemade von Frey hair, about a nickel’s worth of technology.

246 P. Teitelbaum / Behavioural Brain R

Fig. 10. Tracings from photographs taken from the video monitor showing a rightingsequence in which the rat was dropped laterally with respect to the camera. In thefirst photograph (a), the rat is shown immediately upon release so that its head andfeet point skyward. Upon release, the forepaws are tucked up to the sides of the faceand then the head, neck, and shoulders rotate en block toward prone (b and c). Oncethe forequarters are nearly prone, the kindquarters are still 90◦ from prone (d), butthen active pelvic rotation brings the remainder of the body to prone (e). This wash

F

ugbbps

watf

instance, in the study of the regulation of food and water intakein the 1950s and 1960s, we forced in Physiology an entire reap-praisal of the role of taste and appetite, by showing that they were

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;

igh-speed video at 1000 frames/s.

rom Pellis et al. [52].

s to be the best notation system to use, because it is a universaleometrical system for the description of movement, not limitedy having been developed for a particular movement style. Also,ecause EWMN was derived from the analysis of movement in peo-le, it makes sense to study people while trying to computerize theystem.

So, with some sadness, I have closed my animal laboratory. Nowe use movement notation to study the abnormal gait of people

fflicted with Parkinson’s disease. As we work toward achievinghe computerization of this method, we have found two importantacts so far:

esearch 231 (2012) 234– 249

In the deterioration of parkinsonian gait, there is a regression toan infantile form of weight transfer, in which the weight is shiftedforward only after the stepping leg has achieved full contact andsupport by the ground (see Fig. 11, top). In contrast, the normaladult person shifts the weight forward while there is a ground-support in only one leg, as indicated by the release of contact ofthe rear heel from the ground, before the front heel makes contactas the leg steps forward (see Fig. 12) [55]. This is another exampleof reversal of dominance: as in the bandage-backfall, so too in thegait. Tactile contact (support) becomes dominant over vestibularcontrol, so that both legs have to be in contact with the groundbefore weight is shifted forward. (From other aspects of the gate,Forssberg et al. [56] had suggested earlier that parkinsonian gaitresembled 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. How-ever, if we merely ask him to swing his arms when he walks,he instantly automatically walks faster, with longer strides, andwith a more erect posture [55,57] (see Fig. 11, bottom). A similareffect occurs if we merely ask him to take larger steps: he swingshis arms, walks faster, and uses the more adult form of weighttransfer. The principle is: Indirect amplitude modulation via alliedreflexes. In other words, voluntarily augmenting the amplitude ofone of the allied reflexes involved in walking automatically aug-ments the amplitude of the others. Thus, people with Parkinson’sdisease can use these allied reflexes to help themselves to walkbetter [55,57].

20. Force neurophysiology to confront the complexity ofbehavior

“For everything there is a reason, and a time for every matterunder heaven:

. . .. . .. . .. . .. . .

a time to break down, and a time to build up;

. . .. . .. . .. . .. . ..14

The power of Claude Bernard’s method of counterexperimentlies in the fact that he used physical analysis (breaking down) andimmediate resynthesis (building up) in each experiment [3]. Unfor-tunately, science can achieve a great deal of success by continuouslyoversimplifying without an immediate test for resynthesis. As aphysiological psychologist, while studying abnormalities, you canprovide a great service to Neuroscience by forcing an awarenessthat the phenomenon you are studying is a great deal more com-plicated than is presently “conceived of in their philosophy.” For

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.

P. Teitelbaum / Behavioural Brain Research 231 (2012) 234– 249 247

Fig. 11. Top: a person with Parkinson’s disease walking naturally. His arms swingvery little, and he walks slowly, with relatively short steps. The rear heel does notrelease 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. Bot-tom: when asked to swing his arms while walking, he immediately walks faster withlonger steps, and with head more erect. This improves his shift of weight forward,though in this instance, it is still not fully normal.

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 fromthe ground, before the front heel makes contact as the leg steps forward.

essential for regulation in hypothalamic hyperphagia and vital forsurvival after lateral hypothalamic damage. Furthermore, by study-ing recovery rather than concentrating on localization, we showedthat lack of drinking was also critical in the syndrome, and thatmany subtle reflexive, infantile, forms of intake, as in “prandialdrinking” were critical for survival after such brain damage [4].Thus, our work helped to open the field of hypothalamic controlof feeding to psychologists. Now they dominate this field.

An analogous opportunity for physiological psychologists existscurrently in the study of locomotion. In 1981, Cheng and othersin our group [58] showed that after damage in the region of thenucleus reticularis tegmenti pontis (NRTP), a remarkable form ofgalloping locomotion is released in the rat. This form of gallopingis greatly dependent on particular sensory stimuli, thus revealingthe role of many still unknown reflexes in the control of locomo-tion. For instance, the NRTP-damaged rat will gallop headlong intoa wall. The pressure of the wall on the snout instantly shuts downthe galloping, and the rat then sits quietly with its nose in contactwith the wall. In an open field, if a rubber band is twisted around thesnout of such a rat, simulating the wall by applying localized pres-sure on the snout, the rat will remain immobile. If the rubber bandis snipped, however, the rat with such lesions will instantly gallopforward. Therefore, snout contact clearly inhibits forward locomo-

tion (see also the work of Sinnamon [59] on the role of contact inthe inhibition of locomotion).15

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 walkslowly and intermittently. Why? We don’t know.

2 rain R

iNfioNPistsefh

tipigmHtofitwp

2

migtmtoto

sibdtc

2

ostrsp

tbsate

[

[

[

[

[

[

[

[

[

[[

[

[

[

[

[

[

[

[

[

48 P. Teitelbaum / Behavioural B

Haloperidol, which induces akinesia in the normal rat by block-ng dopamine receptors in the brain, does not work at all in theRTP-damaged rat [60]. Nor does morphine. This means that thenal common inhibitory pathway by which dopamine-deficiencyr morphine inhibits locomotion runs through the area of theRTP. This also means that a way of reversing the akinesia ofarkinson’s disease can be achieved by developing techniques fornhibiting the action of the NRTP. Drugs that inhibit the action oferotonin, like methysergide, are effective in doing so [61]. An effec-ive use of serotonin inhibition has not yet been worked out as aupplement to l-dopa in the clinical treatment of Parkinson’s dis-ase, although the effects of serotonin blockade in rats recoveredrom NTRP-damage clearly indicate that such a treatment shouldelp.

This phenomenon is crying out for further analysis and exploita-ion. Maps of the locomotion system make no mention of thenhibitory role of the region of the nucleus reticularis tegmentiontis [62]. Perhaps this is because these findings do not fit

nto neurophysiological thinking about locomotion. This offers areat opportunity for brave young physiological psychologists toake an important contribution to this field [see the work of Vanartesveldt and coworkers [63], and of Sinnamon [59]]. Indeed,

he study of such reflexes was largely abandoned after the deathf Magnus, perhaps due to the invention of more powerful ampli-ers during World War II, so that the young neurophysiologists ofhat time concentrated on microelectrode work instead. There is ahole field of study of compound, allied, reflexes that is waiting forsychologists to breathe life back into it.

1. To obtain funding, build medicine as well as psychology

For a physiological psychologist, building Psychology ofteneans going against the values prevalent among neurologists, and

ndeed, among many medically-oriented physiological psycholo-ists. Since your research funding depends upon their approval,his is critical to your future success. My advice is to do both – build

edicine as well as psychology. Produce important facts abouthe tissue, as well as important facts about the subcomponentsf function. That way you can build Psychology, while continuingo receive approval and financial support from your medically-riented peers.

However, when you talk about your work, I now think youhould not say, “I am working on brain and behavior.” In physiolog-cal psychology, it is too easy to dismiss the role of the analysis ofehavior with wave of the hand, and instead to concentrate on theetails of the mechanics of brain tissue. Say instead, “I am studyinghe relationship of brain to the hierarchy of behavior.” Hierarchyannot be waved away; it must be built.

2. Good research can be great fun

Although it may look to some as though my work has been allver the map, if you think about it, you will see that it has been theame problem all the time (the one that Cliff Morgan mentionedo me, and that Eliot and Alan started out with me on): how to useecovery of function to discover stages in the lateral hypothalamicyndrome (and syndromes related to it) in order to work out therinciples of the hierarchical organization of motivated behavior.

Finally, I should point out that perhaps the most importanthing that I learned from Eliot Steller is that good research cane great fun. Indeed, I have tried to transmit this to my own

tudents by half-jokingly formulating Teitelbaum’s Law: if therere two methods that can be used for doing an experiment (andhere always are), the better method is the one that makes thexperiment more fun to do. This may sound frivolous, but it is

[

[

esearch 231 (2012) 234– 249

not. I have never found it to fail. If you apply this to your ownexperiments, you’ll have tremendous fun. Good luck!

November 29, 1994

Acknowledgements

For helpful criticism, I thank Edward Stricker of the Departmentof Neuroscience at the University of Pittsburgh, Ilan Golani and OferTchernichovsky of the Zoology Department of Tel Aviv University,and Osnat Teitelbaum at the University of Florida.

References

[1] Vallery-Radot R. The life of Pasteur. Garden City, NJ: Doubleday, Page & Com-pany; 1923.

[2] Bernard C. An introduction to the study of experimental medicine. New York:Dover; 1865 [reprinted in English translation, 1957].

[3] Teitelbaum P, Pellis SM. Toward a synthetic physiological psychology. Psycho-logical Science 1992;3:4–20.

[4] Teitelbaum P, Epstein AN. The lateral hypothalamic syndrome: recovery offeeding and drinking after lateral hypothalamic damage. Psychological Review1962;69:74–90.

[5] Descartes R. Philosophical works (1628–1647). New York: Dover; 1955 [trans-lated, Haldane ES, Ross GRT, 1911/12].

[6] Magnus R. Body posture (Korperstellung). Berlin: Springer; 1924.[7] Sherrington CS. The integrative action of the nervous system. New York: Scrib-

ner; 1906.[8] Tinbergen N. The study of instinct. Oxford: Clarendon Press; 1951.[9] Lorenz KZ. The foundations of ethology. New York: Springer Verlag; 1981.10] Pavlov IP. Conditioned reflexes. New York: Dover; 1927 [reprinted in this

English translation, 1960].11] Skinner BF. The behavior of organisms: an experimental analysis. New York:

Appleton-Century; 1938.12] Teitelbaum P. The lateral hypothalamic double-disconnection syndrome: a

reappraisal and a new theory of recovery of function. In: Hulse SH, Green BF,editors. One hundred years of psychological research in America: G. StanleyHall and the Johns Hopkins tradition. Baltimore: The Johns Hopkins UniversityPress; 1986. p. 79–124.

13] Hetherington AW, Ranson SW. Hypothalamic lesions and adiposity in the rat.Anatomical Record 1940;78:149–72.

14] Brobeck JR, Tepperman L, Long CNH. Experimental hypothalamic hyperphagiain the albino rat. Yale Journal of Biology and Medicine 1943;15:831–5.

15] Stellar E, Krause NP. New stereotaxic instrument for use with the rat. Science1954;120:664–6.

16] Teitelbaum P, Stellar E. Recovery from failure to eat produced by hypothalamiclesions. Science 1954;120:894–5.

17] Anand BK, Brobeck JR. Localization of a feeding center in the hypothalamusof the rat. Proceedings of the Society of Experimental Biology and Medicine1951;77:323–4.

18] Anand BK, Brobeck JR. Hypothalamic control of food intake. Yale Journal ofBiology and Medicine 1951;24:123–40.

19] Stellar E. The physiology of motivation. Psychological Review 1954;61:5–22.20] Teitelbaum P, Stricker EM. Compound complementarities in the study of moti-

vated behavior. Psychological Review 1994;101:312–7.21] Teitelbaum P. Sensory control of hypothalamic hyperphagia. Journal of Com-

parative and Physiological Psychology 1955;48:156–63.22] Sclafani A, Springer D, Kluge L. Effects of quinine-adulterated diets on the food

intake and body weight of obese and non-obese hypothalamic hyperphagicrats. Physiology and Behavior 1976;16:631–40.

23] Teitelbaum P. Appetite. Proceedings of the American Philosophical Society1964:464–72, 108.

24] Williams DR, Teitelbaum P. Some observations on the starvation resulting fromhypothalamic hyperphagia in the albino rat. Journal of Comparative and Phys-iological Psychology 1959;52:458–65.

25] Faber KH. Nosography: the evolution of clinical medicine in modern times. NewYork: Paul H. Hoeber, Inc.; 1930 [reprinted, New York: AMS Press; 1978].

26] Bolles RC. Why you should avoid statistics. Biological Psychaitry1988;23:79–85.

27] Whishaw IQ. The decorticate rat. In: Kolb B, Tees RC, editors. The cerebral cortexof the rat. Cambridge, MA: MIT Press; 1990. p. 239–67.

28] Teitelbaum P, Cheng M-F, Rozin P. Development of feeding parallels its recoveryafter hypothalamic damage. Journal of Comparative and Physiological Psychol-ogy 1969;67:430–41.

29] Cheng M-F, Rozin P, Teitelbaum P. Starvation retards the development of food

and water regulations. Journal of Comparative and Physiological Psychology1971;76:206–18.

30] Kintsch W, Cacioppo JT. Introduction to the 100th anniversary issue of ThePsychological Review. Psychological Review 1994;101:195–9.

31] Dewsbury DA. Psychobiology. American Psychologist 1991;46:198–205.

rain R

[

[

[

[

[

[

[

[

[

[

[

[

[

[

[

[

[

[

[

[

[

[[

[

[

[

[

[

[

[

P. Teitelbaum / Behavioural B

32] Holton G. Thematic origins of scientific thought. Kepler to Einstein. Cambridge,MA: Harvard University Press; 1973.

33] Teitelbaum P, Pellis VC, Pellis SM. Can allied reflexes promote the integration ofa robot’s behavior? In: Meyer J-A, Wilson SW, editors. From animals to animats:simulation of adaptive behavior. Cambridge, MA: MIT Press; 1991. p. 97–104.

34] Hubbard R, Wald G. The mechanism of rhodopsin synthesis. Proceedings of theNational Academy of Sciences of the United States of America 1951;37:69–79.

35] Marshall JF, Turner BH, Teitelbaum P. Sensory neglect produced by lateralhypothalamic damage. Science 1971;174:523–5.

36] Marshall JF, Teitelbaum P. Further analysis of sensory inattention followinglateral hypothalamic damage in rats. Journal of Comparative and PhysiologicalPsychology 1974;86:375–95.

37] O’Brien DP, Chesire RM, Teitelbaum P. Vestibular versus tail-pinch-inducedactivation in cats with lateral hypothalamic lesions. Physiology and Behavior1985;34:811–4.

38] Szechtman H. Redirected oral behavior in rats induced by tail pinch and elec-trical stimulation of the tail. Physiology and Behavior 1980;24:57–64.

39] Wolgin DL, Teitelbaum P. Role of activation and sensory stimuli in recoveryfrom lateral hypothalamic damage in the cat. Journal of Comparative and Phys-iological Psychology 1978;92:474–500.

40] Beaumont W, Osler W. Experiments and observations on the gastric juice, andthe physiology of digestion. New York: Dover Publications; 1833 [reprinted inthis edition, 1959].

41] Van Harreveld A, Bogen JE. The clinging position of the bulbocapninized cat.Experimental Neurology 1961;4:241–61.

42] Teitelbaum P, Schallert T, DeRyck M, Whishaw IQ, Golani I. Motor subsystemsin motivated behavior. In: Thompson RF, Hicks LH, Shvyrkov VB, editors. Neu-ral mechanisms of goal-directed behavior and learning. New York: AcademicPress; 1980. p. 127–43.

43] Schallert T, Whishaw IQ, DeRyck M, Teitelbaum P. The postures ofcatecholamine-depletion catalepsy: their possible adaptive value in ther-moregulation. Physiology and Behavior 1978;21:817–20.

44] Schallert T, DeRyck M, Whishaw IQ, Ramirez WD, Teitelbaum P. Excessive brac-ing reactions and their control by atropine and l-dopa in an animal analog ofparkinsonism. Experimental Neurology 1979;64:33–43.

45] DeRyck M, Schallert T, Teitelbaum P. Morphine versus haloperidol catalepsy inthe rat: a behavioral analysis of postural support mechanisms. Brain Research1980;201:143–72.

46] Morrissey TK, Pellis SM, Pellis VC, Teitelbaum P. Seemingly paradoxical jump-

ing is triggered by postural instability in cataleptic haloperidol-treated rats.Behavioural Brain Research 1989;35:195–207.

47] Teitelbaum P. The physiological analysis of motivated behavior. In: ZimbardoPG, Ruch FL, editors. Psychology and life. 9th ed. Glenview, IL: Scott, Foresman& Co.; 1977. p. 2A-F.

[

[

esearch 231 (2012) 234– 249 249

48] Teitelbaum P, Wolgin DL, DeRyck M, Marin OSM. Bandage-backfall reaction:occurs in infancy, hypothalamic damage and catalepsy. Proceedings of theNational Academy of Sciences of the United States of America 1976;73:3311–4.

49] Cordover AJ, Pellis SM, Teitelbaum P. Haloperidol exaggerates proprioceptive-tactile support reflexes and diminishes vestibular dominance over them.Behavioural Brain Research 1993;56:197–201.

50] Levitt D, Somnolence Teitelbaum P. akinesia, and sensory activation ofmotivated behavior in the lateral hypothalamic syndrome. Proceedingsof the National Academy of Sciences of the United States of America1975;72:2819–23.

51] Golani I, Wolgin DL, Teitelbaum P. A proposed natural geometry of recoveryfrom akinesia in the lateral hypothalamic rat. Brain Research 1979;164:237–67.

52] Pellis SM, Pellis VC, Teitelbaum P. Air-righting without the cervical neck-righting reflex in rats. Behavioural Brain Research 1991;46:185–8.

53] Feldenkrais M. The elusive obvious. Cupertino, CA: Meta Publications; 1981.54] Eshkol N, Wachman A. Movement notation. London, UK: Weidenfeld and

Nicholson; 1958.55] Teitelbaum P, Maurer RM, Fryman J, Teitelbaum OB, Vilensky J, Creedon MP.

Dimensions of disintegration in the stereotyped locomotion characteristic ofparkinsonism and autism. In: Sprague RL, Newell KM, editors. Stereotypedmovements: brain–behavior relationships. Washington, DC: American Psycho-logical Association; 1996. p. 167–93.

56] Forrsberg HB, Johnels B, Steg G. Is parkinsonian gait caused by a regression toan immature walking pattern. Advances in Neurology 1984;40:375–9.

57] Behrman A, Teitelbaum P, Cauraugh JH. Verbal instructional sets normalize thetemporal and spatial gait variables in Parkinson’s disease. Journal of Neurology,Neurosurgery & Psychiatry 1998;65:580–2.

58] Cheng J-T, Schallert T, DeRyck M, Teitelbaum P. Galloping induced by pontinetegmentum damage in rats: a form of Parkinsonian festination not blocked byhaloperidol. Proceedings of the National Academy of Sciences of the UnitedStates of America 1981;78:3279–83.

59] Sinnamon HM. Preoptic and hypothalamic neurons and the initiation of loco-motion in the anesthetized rat. Progress in Neurobiology 1993;41:323–44.

60] Chesire RM, Cheng J-T, Teitelbaum P. The inhibition of movement by mor-phine or haloperidol depends on an intact nucleus reticularis tegmenti pontis.Physiology and Behavior 1983;30:809–18.

61] Chesire RM, Cheng J-T, Teitelbaum P. Reinstatement of festinating forward loco-motion by antiserotonergic drugs in rats partially recovered from damage inthe region of the nucleus reticularis tegmenti pontis. Experimental Neurology

1982;77:286–94.

62] Garcia-Rill E. The basal ganglia and the locomotor regions. Brain Research1986;396:47–63.

63] Van Hartesveldt C, Sickles AE, Porter JD, Stehouwer DJ. l-Dopa-induced air-stepping in developing rats. Developmental Brain Research 1991;58:251–5.