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Effects of peptides on animal and human behavior: a review of studies published in the first twenty years of the journal Peptidesa
Peptides 22 (2001) 2181–2255
Review
Effects of peptides on animal and human behavior: A review of studies published in the first twenty years of the journal Peptides Robert N. McLaya, Weihong Panb,*, Abba J. Kastinb a
b
Naval Medical Center San Diego, Department of Psychiatry, San Diego, CA, USA VA Medical Center and Tulane University School of Medicine, Neuroscience Training Program and Department of Medicine, New Orleans LA 70112-1262, USA Received June 25, 2001; accepted July 27, 2001
Abstract This review catalogs effects of peptides on various aspects of animal and human behavior as published in the journal Peptides in its first twenty years. Topics covered include: activity levels, addiction behavior, ingestive behaviors, learning and memory-based behaviors, nociceptive behaviors, social and sexual behavior, and stereotyped and other behaviors. There are separate tables for these behaviors and a short introduction for each section. © 2001 Elsevier Science Inc. All rights reserved. Keywords: Peptides; Behavior; Activity; Addiction; Ingestion; Learning; Memory; Nociception; Sex; Stereotypy
1. Introduction Why do we do the things we do? This question could be answered on many levels, but increasingly research is discovering ways in which behavior is affected, and sometimes governed, by the actions of peptides. For over twenty years this journal has published articles concerning peptides, many of which involved effects on animal and human behavior. This review is an attempt to catalog those findings. We have focused on studies in which a behavior was elicited after the administration of a peptide. Other types of
studies, such as those studying animals deficient in a particular peptide, or using an agonist to block the effect of a peptide, are for the most part not included, except where the antagonist itself is a peptide or protein. We by no means discount the importance of such studies in understanding how peptides influence behavior, however. Readers wishing to find listings of other effects may be served by reviews previously published in Peptides [3,412,607]. In choosing the articles to include, decisions had to be made concerning what constitutes animal behavior. Usually the choice was obvious. A study of classically conditioned
* Corresponding author. Tel.: ⫹1-504-589-5928; fax: ⫹1-504-522-8559. E-mail address: [email protected] (W. Pan). Abbreviations: 5-HT, Serotonin; 5-HTP, 5-Hydroxytryptophan; 6-OHDA, 6-Hydroxydopamine; A18fa, Ala-Gly Glu-Gly Leu-Ser-Ser-ProPhe-Trp-Ser-Leu-Ala-Ala-Pro-Gln-Arg-Phe-NH2, Admin., Administration; Act., Activity; ACTH, Adrenocorticotropic Hormone; ANF, Atrial Natriuretic Factor; AT, Angiotensin; BBS, Bombesin; BUBUC, Tyr-DCys(StBu)-Gly Phe-Leu-Thr-(OtBu); CCK, Cholecystokinin; CCK-4, Cholecystokinin Tetrapeptide; Tetragastrin; CCK-8, Cholecystokinin Octapeptide; Sincalide; CGRP, Calcitonin Gene-Related Peptide; CRF, Corticotropin Releasing Factor; CTAP, D-Phe-Cys-Try-D-Trp-ORn-Thr-PenThr-NH2; CTOP, D-Phe-Cys-Tyr-D-Tyr-Orn-Thr-NH2; DADELT II, [D-Ala2]Deltorphin; DADLE, D-Ala2-D-Leu5-Enkephalin; DAGO, D-Ala2-MePhe4-Gly-ol5-Enkephalin; DALA, D-Ala2-Methionine-enkephalinamide; DALCE, [D-Ala2,Leu5,Cys6]-Enkephalin; DAMA, D-Ala2Met-Enkephalin Amide; DAME, [D-Ala2-Met5]Enkephalinamide; DAMCK, Tyr-D-Ala-Gly-(NMe)Phe-CH2Cl; DAMGO, Tyr-D-Ala-Gly-
(NMe)Phe-Gly-ol; DDAVP, Desmopressin; 1-deamino-Cys-8-D-Arg-Vasopressin; DGAVP; Desglycinamide-arginine8-Vasopressin; DPEN [D-Pen2,D-Pen5]-Enkephalin; DPDPE, [D-Pen2,D-Pen5]-Enkephalin; DSIP, Delta Sleep-Inducing Peptide; EGF, Epidermal Growth Factor; F8Fa, Phe-Leu-Phe-Gln-Pro-Gln-Arg-Phe-NH2; Frag., Fragment; FMRFa, Phe-Met-Arg-Phe-amide; GHRH, Growth Hormone-Releasing Hormone; GR73632, D-Ala1-[L-Pro9,Me-Leu8]Substance P-(7–11); Ind., Induced; LHRH, Luteinizing Hormone Releasing Hormone; LLTNAM, Lys (CH2NH)-Trp(Nps)-OMe; MCH, Melanin Concentrating Hormone; MIF-1, Melanotropin (MSH) Release-Inhibiting Factor-1; MSH, Melanocyte Stimulating Hormone; NT, Neurotensin; NP-, Neuropeptide-; OT, Oxytocin; Org2766, H-Met-(O2)-Glu-His-Phe-D-Lys-Phe-OH; PACAP, Pituitary Adenylate Cyclase-Activating Polypeptide; PG-KII, pGlu-ProAsn-Pro-Asp-Glu-Phe-Val-Gly-Leu-Met-NH2; PVN, Paraventricular Nucleus; SP, Substance P; TRH, Thyrotropin-Releasing Hormone; TyrMIF-1, Tyr-Pro-Leu-Gly-NH2; Tyr-W-MIF-1, Tyr-Pro-Trp-Gly NH2; VP, Vasopressin; VIP, Vasoactive Intestinal Peptide; YPLG, Tyr-Pro-Leu-Gly
0196-9781/01/$ – see front matter © 2001 Elsevier Science Inc. All rights reserved. PII: S 0 1 9 6 - 9 7 8 1 ( 0 1 ) 0 0 5 5 0 - 2
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eye-blink response is clearly a behavioral study. A study of body temperature is not. In the case of classically conditioned temperature response [65], the choice is less clear. In general, we limited the included studies to those of externally observable actions of an animal or person. Where a study clearly added to the understanding of behavior, however, we were liberal in inclusion. In some cases this may seem somewhat arbitrary. Made only slightly less arbitrary by the existence of authors’ keywords is the decision on how to categorize behavior. We have divided behaviors into seven, broad groups: activity levels, addiction behaviors, ingestive behaviors, learning and memory-based behaviors, nociceptive behaviors, social and sexual behaviors, and stereotyped and other behaviors. We realize that there are many areas of overlap and many ways by which an individual might wish the information organized. Readers with access to the internet and a desire to reorganize the tables to their own liking can find electronic versions of the tables at: http://www.sam.tulane.edu/ classware/pathology/medical_pathology/mcpath/peptides behavior.html For those whose access is limited to the printed page, we hope that these categories are useful. All peptides were administered peripherally unless otherwise noted.
2. Topics 2.1. Activity level Any study of behavior is dependent on the activity level of the organism. Simply put, an animal that is hyperstimulated gives a different response than one that is unconscious or paralyzed. Because of this, motor activity, as measured by distance moved in an enclosed space, is often one of the first effects noted for a peptide. Despite the wealth of data, there is no unifying mechanism yet known to underlie activity level. Some peptides appear to work though common neural substrates such as the ventral striatum and globus pallidus to produce increased activity, whereas others may be independent of such pathways [530]. Still others may regulate activity without directly acting on the brain at all, but rather through peripheral control of metabolism [270]. Peptides appear to be particularly important in the regulation of daily activity and sleep cycles. Balance between neuropeptides such as GHRH and CRF is a key modulator of wakefulness [516]. Other substances such as melatonin are not only believed to be endogenous regulators of sleep, but also have advanced to the clinical stage as pharmaceuticals [192].
Table 1 lists effects on activity published in Peptides. 2.2. Addictive behavior Peptides are critical to the neurobiology of addiction. Often they are the endogenous substrates for the receptors on which drugs of abuse act. The endogenous peptides and their analogs thus have great potential for antagonizing the effects of drugs, and for producing safer, less-addictive versions of addictive substances. Perhaps more importantly, peptide neurotransmitters may underlie certain, central mechanisms of addiction. Drugs of addiction act at a variety of different brain areas and receptors. Common mechanisms of addiction appear to come into play with multiple drugs, however. The mesolimbic dopamine system is the best known [22], but other, peptide-mediated systems also are critical to addiction. Opioid peptide systems within and independent of the mesolimbic system play a role in drug-use reinforcement. CRF and other stress-peptides are critical in withdrawal behaviors, and other peptide neurotransmitters may modulate transmission within the dopamine system [289]. In short, the system is hooked on peptides. Table 2 lists effects on addictive behaviour. 2.3. Nociceptive behavior The mounting of an appropriate response to pain depends on a variety of mechanisms. The painful stimuli must be relayed to the central nervous system. That stimuli must then be appropriately received and a response formed. The signal for the response is then sent back from the nervous system, and the end organ must be capable of carrying out the appropriate action. In some cases pain behaviors can be elicited despite an absence of one or more of these steps. For example, a person might withdraw his hand from an incoming needle because of an expectation of pain even though no pain is or will be experienced. For this review we have relegated such complex behaviors to the learning and memory-related behaviors section. A focus on simple and immediate actions of pain, however, still leaves a number of areas in which peptides influence behavior. The first detector of pain is often referred to as a nociceptor. However, no specific histologic or molecular structure can specifically be said to constitute that receptor. It is believed that noxious stimuli are initially detected by irritation of free, unmyelinated terminal arborizations of nerves. Nociceptive messages are then transmitted by thin myelinated (A-␦) or non-myelinated (C) fibers. A number of substances, including bradykinin, bombesin, galanin, somatostatin, cholecystokinin, vasoactive intestinal peptide, neuropeptide Y, capsaicin, opioids, and cytokines, influence detection and transmission in these nociceptive fibers [43]. Nociceptive signals are carried in the spinal cord via segmental and supraspinal mechanisms. The dorsal horn of
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
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Table 1 Activity levels Peptide
Movement
Species
Effect
Notes
Vol
Page
Ref
ACTH
Exploratory behavior
Rat
Increased
5
713
546
ACTH ACTH
Motor act. Motor act.
Rat Rat
4 3
117 117
48 48
ACTH frag. (18–24) ACTH frag. (18–24) ACTH frag. (20–24) ACTH/MSH (4–10) analog ebiratide
Sleep Sleep Sleep Sleep
Rat Rat Rat Human
Decreased Increase followed by decrease Increased No effect Increased Decreased
Pre-optic, anterior hypothalamic admin. Chronic admin. Not naltrexone reversible ICV; paradoxical sleep ICV; paradoxical sleep ICV; paradoxical sleep Increased sleep onset latency and increased wakefulness
15 15 15 12
237 237 237 1007
582 582 582 515
ACTH/MSH 4–10 analog ORG 2766 ACTH/MSH 4–10 analog ORG 2766 ACTH/MSH 4–10 analog ORG 2766
Motor act.
Rat
No change
2
255
174
Motor act.
Rat
Increased
First three weeks of life
6
2S 41
1
Motor act.
Rat
Decreased
13
541
235
ACTH/MSH 4–10 analog ORG 2766
Rearing
Rat
Increased
Amygdala admin.; in isolated animals but not controls Rearing and grasping ability occurs earlier in development
6
2S 41
1
ACTH/MSH 4–10 analog ORG 2766 ACTH/MSH frag. (4–10) ACTH/MSH frag. (4–10)
Rearing
Rat
Decreased
5
1061
203
Motor act. Motor act.
Rabbit Rat
No change Increased
6 6
2S 97 285
229 189
ACTH/MSH frag. (4–10) ACTH/MSH frag. (4–10) ACTH/MSH frag. (4–10)
Motor act. Motor act. Motor act.
Rat Rat Rat
Increased Increased Decreased
6 7 9
2S 41 201 277
1 360 593
ACTH/MSH frag. (4–10)
Motor act.
Rat
Increased
9
151
458
ACTH/MSH frag. (4–10)
Rearing
Rat
Increased
6
2S 41
1
Adrenal Peptide E (Frog) Amylin
Motor act. Motor act.
Mouse Rat
No change Decreased
17 17
1291 589
97 91
AT II
Motor act.
Rat
Decreased
9
475
58
BBS BBS BBS
Exploratory behavior Exploratory behavior Motor act.
Rat Rat Rat
Increased Decreased Increased
4 12 4
405 761 693
103 182 355
BBS
Motor act.
Rat
Increased
9
1S 245
260
BBS
Rearing
Rat
Increased
4
693
355
BBS
Sleep
Rat
Decreased
8
237
303
BUBUC
Rearing
Mouse
Increased
11
983
188
In stressed, hypophysectomized rats First three weeks of life In stressed but not controls Neonates; age dependent effect Rearing and grasping ability occurs earlier in development ICV ICV; peptide also antagonized amphetamine-induced hyperactivity; effect blocked by ritanserin ICV; in combination with saralasin but not alone ICV ICV; effect antagonized by neuroleptics With admin. to nucleus accumbens but not nucleus tractus solitarius ICV; effect antagonized by neuroleptics Admin. to multiple brain areas Antagonized by naltrindole
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Table 1 (continued) Peptide
Movement
Species
Effect
Notes
Vol
Page
Ref
Casomorphin ()
Sleep
Rat
Neonates; effect blocked by naloxone
11
1
533
Casomorphin analog BCH 325
Motor act.
Rat
Increased quiet sleep decreased active sleep Decreased
Apomorphine-induced hypoactivity; antagonized by sulpiride
16
635
447
Casomorphin analog BCH 325
Sleep
Rat
12
893
483
Casomorphin derivative BCH 325 CCK
Motor act.
Rat
Decreased at low dose, increased at high dose Increased
13
141
287
Exploratory behavior
Rat
Decreased
4
743
109
CCK
Motor act.
Rat
Decreased
7
587
455
CCK analog Boc[diNle]CCK-7 CCK analog Boc[Phe(2,6diMe)7]-CCK-7 CCK analog Boc-[Phe(2Me)7]-CCK-7 CCK analog Boc-[Phe(4Me)7]-CCK-7 CCK analog Boc[Phe(pentaMe)7]-CCK-7 CCK analog Boc[Phe(triMe)7]-CCK-7 CCK analog Boc-CCK-7 CCK-4 CCK-4
Motor act.
Rat
Decreased
In stress-induced hypoactivity Effect blocked by lesions of nucleus tractus solitarius Effect blocked by vagotomy or capsaicin admin. CNS admin.
6
415
461
Rearing
Mouse
Decreased
ICV
19
301
333
Rearing
Mouse
Decreased
ICV
19
301
333
Rearing
Mouse
Decreased
ICV
19
301
333
Rearing
Mouse
No change
ICV
19
301
333
Rearing
Mouse
Decreased
ICV
19
301
333
Rearing Circling Exploratory behavior
Mouse Rat Rat
Decreased No change Decreased
19 7 16
301 809 815
333 367 215
CCK-4
Motor act.
Rat
Decreased
6
91
275
CCK-4
Rearing
Rat
Decreased
6
91
275
CCK-8 CCK-8
Circling Exploratory behavior
Rat Rat
Increased Decreased
7 5
809 529
367 272
CCK-8
Motor act.
Mouse
Decreased
13
155
231
CCK-8
Motor act.
Rat
No change
4
749
480
CCK-8 CCK-8
Motor act. Motor act.
Rat Rat
Decreased Decreased
6 6
53 91
254 275
CCK-8 CCK-8
Rearing Rearing
Mouse Rat
Decreased Decreased
19 4
301 749
333 480
CCK-8
Rearing
Rat
Decreased
ICV ICV In elevated plus maze, but not in open field. With admin. to nucleus accumbens but not other brain areas With admin. to nucleus accumbens but not other brain areas ICV ICV or IV; antagonized by proglumide ICV or IP; antagonized by CCK-A, but not CCK-B, antagonists Ventral tegmental area and nuclei accumbens septi admin. ICV; in VIP-treated rats With admin. to nucleus accumbens but not other brain areas ICV Ventral tegmental area and nuclei accumbens septi admin. With admin. to nucleus accumbens but not other brain areas
6
91
275
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Table 1 (continued) Peptide
Movement
Species
Effect
Notes
Vol
Page
Ref
CCK-8
Sleep
Mouse
Increased
3
701
610
CCK-8, desulfated Ceruletide analogs Ceruletide analogs
Circling Exploratory behavior Sleep
Rat Mouse Mouse
No change Decreased Increased
Hexobarbital-induced sleep ICV
7 2 3
809 2S 701
367 609 610
CGRP
Motor act.
Rat
Increased
17
1183
294
Corticotropin-like intermediate lobe peptide CRF CRF CRF
Sleep
Rat
Increased
15
237
582
Exploratory behavior Motor act. Motor act.
Monkey Rat Rat
Increased Increased Increased
4 6 6
211 3S 53 923
267 392 161
CRF
Motor act.
Rat
Increased
7
597
476
D-Arg dipeptides
Motor act.
Mouse
Decreased
6
35
475
D-Arg tetrapeptides
Motor act.
Mouse
6
35
475
Dermorphin
Sleep
Chicken
Decrease followed by increase Increased
6
3S 137
397
DSIP
Motor act.
Mouse
Decreased
3
729
211
(1–13) (1–6) (1–7) (1–8)
Motor act. Rearing Circling Circling Circling
Rat Rat and mouse Rat Rat Rat
Biphasic effect No change Increased Increased Increased
3 1 8 8 8
623 341 837 837 837
210 578 186 186 186
Dynorphin frag. (2–17) EGF Endorphin (␣) Endorphin (␣) Endorphin (␣)
Circling Motor act. Motor act. Motor act. Motor act.
Rat Sheep Mouse Mouse Mouse
Increased Decreased No change Decreased Decreased
8 9 13 5 5
837 1221 737 129 1011
186 417 271 555 554
Endorphin (␣) Endorphin ()
Rearing Motor act.
Mouse Goldfish
Decreased Increased
5 3
129 679
555 277
Endorphin () Endorphin () Endorphin ()
Motor act. Motor act. Motor act.
Mouse Rat Rat
No change Decreased Increased
13 1 1
737 179 179
271 603 603
Endorphin () Endorphin () Endorphin ()
Motor act. Motor act. Sleep
Rat Sheep Chicken
Increased Decreased Increased
6 9 6
189 1221 3S 137
601 417 397
Enkephalin (Leu) Enkephalin analog DPDPE Enkephalin analog DPDPE
Circling Motor act.
Rat Rat
Increased Increased
8 13
837 755
186 509
Motor act.
Rat
Increased
14
511
509
DSIP Dynorphin Dynorphin Dynorphin Dynorphin
frag. frag. frag. frag.
Hexobarbital-induced sleep Antagonized by nitric oxide synthase inhibitor ICV; paradoxical sleep
With ICV but not SC ICV; greater effect in controls than in hypophysectomized ICV; blocked by naloxone ICV; mildly antagonized by naloxone ICV; hyperactivity antagonized by naloxone ICV; chicks; effect inhibited by naloxone In amphetamine-treated mice; only at selected doses ICV Substantia nigra admin. Substantia nigra admin. Substantia nigra admin.; not antagonized by naloxone Substantia nigra admin. ICV ICV ICV; not antagonized by naloxone At constant temperatures, but not with temp. gradient
In amphetamine-treated, but not controls ICV ICV; chicks; effect inhibited by naloxone Substantia nigra admin. ICV; morphine-induced hyperactivity ICV; morphine-treated rats
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Table 1 (continued) Peptide
Movement
Species
Effect
Notes
Vol
Page
Ref
analog
Sleep
Chicken
Increased
6
3S 137
397
analog
Motor act.
Rat
Decreased
14
511
509
analog
Motor act.
Rat
Decreased
1
139
257
analog
Motor act.
Human
No change
ICV; chicks; effect inhibited by naloxone ICV; DPDPE and morphine-treated rats ICV; antagonized by taurine but not GABA Elderly
12
871
75
Motor act.
Rat
No change
10
403
443
Mouse
No change
16
1283
556
Galanin
Alternating movements Motor act.
Antagonized morphineinduced horizontal, but not vertical, movement ICV
Rat
No change
11
995
304
Galanin Gastrin I (1–17) GH GHRH GHRH Growth hormone
Motor Motor Motor Motor Motor Motor
act. act. act. act. act. act.
Rat Rat Rat Rat Rat Rat
No change No change Decreased Increased Increased Decreased
Periventricular and PVN admin. PVN admin.
13 1 14 14 14 14
323 383 707 707 708 708
354 408 7 7 7 7
Kassinin-like peptide TK PG-KII L-pyroglutamyl-Lleucine MIF-1
Motor act.
Rat
Increased
18
825
437
Jumping
Rat
Delayed
4
417
295
Circling
Rat
No change
2
189
148
MIF-1
Motor act.
Mouse
1
293
147
MIF-1
Motor act.
Rat
Potentiated increase and suppressed decrease No change
Morphine-withdrawalinduced jumping ICV; dopamine or 5HT stimulated circling Morphine-treated
At 4° or 20°C; not altered by melanin or hypophysectomy
1
243
600
MIF-1 MIF-1
Motor act. Motor act.
Rat Rat
Increased Increased
6 12
189 915
601 293
MIF-1
Motor act.
Mouse
1
293
147
MIF-1 Motilin MSH (␣)
Sleep Running Exploratory behavior
Rat Rat Rat
Potentiated increase and suppressed decrease Increased No change Decreased
6 1 17
7 383 171
440 408 207
MSH (␣)
Motor act.
Rat
No change
1
243
600
MSH (␣) MSH (␣)
Motor act. Motor act.
Rat Rat
No change Increased
2 4
255 865
174 598
MSH (␣) MSH (␣)
Motor act. Motor act.
Rat Rat
Increased Increased
6 16
189 821
601 110
MSH (␣)
Rearing
Rat
Increased
16
821
110
Enkephalin DALA Enkephalin DALCE Enkephalin DAMA Enkephalin DDAVP FMRFa
Galanin
Inhibits habituation Effect with acute but not chronic, treatment ICV
In forced swim test; synergy with tricyclic antidepressants Morphine-induced motor act.
Alcohol-induced sleep Admin. to medial preoptic area; antagonized by MCH At 4° or 20°C; not altered by melanin or hypophysectomy Amphetamine-treated; interactions with L/D cycle and age ICV; antagonized by diazepam or baclofen ICV; antagonized by diazepam or baclofen
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Table 1 (continued) Peptide
Movement
Species
Effect
N-acetylmuramyl-Lalanyl-D-isoglutamine
Sleep (REM)
Rat
Naltrindole
Motor act.
Rat
Decrease followed by increase (no overall, average change) No change
NP-E-I
Motor act.
Rat
Increased
NP-E-I
Rearing
Rat
Increased
NPY NPY NPY NPY
Motor act. Motor act. Resting Sleep cycles
Rat Rat Hamster Hamster
Decreased Decreased Decreased Phase shift
NPY antibody
Circling
Mouse
Increased
NPY antibody
Motor act.
Mouse
Increased
NPY frag. (1–30) NT
Motor act. Motor act.
Rat Guinea pig
Decreased Increased
NT NT NT
Motor act. Motor act. Motor act.
Mouse Mouse Mouse
Increased Decreased Decreased
NT NT NT
Motor act. Motor act. Motor act.
Rat Rat Rat
NT
Motor act.
Rat
No change Decreased Complex interactions with dose and enkephalinase inhibitors Decreased
NT NT NT analog [D-Phe11]-NT NT analog [D-Tyr11]-NT NT analog [Phe11]-NT NT analog Eisai hexapeptide
Rearing Sleep Motor act. Motor act. Motor act. Motor act.
Rat Rat Rat Rat Rat Mouse
No change No change Increased Increased Decreased Decreased
Omega-agatoxin IVA
Motor act.
Rat
No change
Omega-conotoxin GVIA
Motor act.
Rat
No change
Opiate antagonist WIN 44,441–3
Motor recovery
Cat
Increased
Notes
ICV; DPDPE- and morphine-treated rats ICV; antagonized by MSH ICV; antagonized by MSH ICV ICV ICV Geniculohypothalamic tract admin.; animals housed in darkness With admin. to ventromedial thalamus but not other brain areas With admin. to hypothalamus but not other brain areas ICV Acted as noxious stimulus Intrathecal admin. Intrathecal admin. Amphetamine- or phencyclidine-induced hyperactivity PVN and IV admin. ICV
ICV; not antagonized by non-peptide NT antagonist PVN and IV admin. PVN and IV admin.
Amphetamine- or phencyclidine-induced hyperactivity ICV; peptides antagonized druginduced hyperactivity ICV; peptides antagonized druginduced hyperactivity Spinal cord injury
Vol
Page
Ref
5
91
184
14
511
509
18
393
470
18
393
470
15 14 9 15
799 909 1389 1475
54 428 300 245
15
607
579
15
607
579
15 13
799 841
54 454
4 4 18
517 517 1223
249 249 473
4 2 11
493 171 551
511 261 399
16
37
442
4 4 2 2 2 18
493 493 171 171 171 1223
511 511 261 261 261 473
19
1017
404
19
1017
404
6
1S 15
169
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Table 1 (continued) Peptide
Movement
Species
Effect
Notes
Vol
Page
Ref
OT
Motor act.
Mouse
Increased
5
535
351
OT Saralasin
Sleep Motor act.
Rat Rat
Increased Decreased
ICV; effect enhanced by co-admin. of amastatin Alcohol-induced sleep ICV; in combination with AT II, but not alone
6 9
7 475
440 58
Sauvagine Sauvagine Sauvagine
Motor act. Motor act. Resting
Mouse Rat Rat
Decreased Increased Increased
6 6 6
3S 59 3S 53 3S 53
420 392 392
Secretin Secretin SK
Motor act. Rearing Motor act.
Rat Rat Rat
Decreased No change Increased
4 4 6
739 739 2S 113
83 83 144
Somatostatin Somatostatin frag. (3–8), (9–14), or (7–10) SP
Rearing Rearing
Rat Rat
Decreased No change
10 10
1153 1153
569 569
Motor act.
Rat
Increased
1
103
262
SP SP
Motor act. Motor act.
Rat Rat
No change Increased
9 6
937 2S 113
173 144
SP frag. (1–7) SP frag. (1–7)
Motor act. Vertical movement
Mouse Rat
Increased Increased
4 20
763 1437
219 126
SP frag. analog pyroglutamylSP(7–11) TRH TRH TRH
Motor act.
Mouse
Decreased
4
763
219
Motor act. Motor act. Motor act.
Rat Rat Rat
Decreased Increased Prevented immobility
2 4 5
131 239 743
602 155 403
TRH
Motor act.
Rat
Increased
6
851
605
TRH
Motor act.
Rat
Increased
5
899
276
TRH analog CG 3704
Motor act.
Rat
Increased
6
851
605
TRH analog DN-1417
Motor act.
Rat
Prevented immobility
5
743
403
TRH analog DN-1417 TRH analog MK-711
Motor act. Motor act.
Rat Rat
Increased Prevented immobility
6 5
1093 743
386 403
TRH analog PS-24 TRH analog RX77386 Urotensin I Urotensin I
Motor act. Motor act. Motor act. Resting
Rat Rat Rat Rat
Increased Increased Increased Increased
2 11 6 6
131 897 3S 53 3S 53
602 258 392 392
Vasotocin Vasotocin
Motor act. Sleep
Cat Cat
Decreased Increased
5 5
25 25
204 204
With ICV but not SC Compared with fasting controls ICV ICV Ventral tegmental area admin. ICV ICV ICV; in controls and haloperidol-treated, but not FLA-63- or methyltyrosine-treated ICV Ventral tegmental area admin. Dorsal periaqueductal gray matter admin.
ICV Depression model; effect present in and after swim, but not before No effect of iron deficiency ICV; effect enhanced by ablation of frontal cortex No effect of iron deficiency Depression model; effect present in and after swim, but not before Depression model; effect present in and after swim, but not before Neonates With ICV but not SC Compared to fasting controls Neonates Neonates
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2189
Table 1 (continued) Peptide
Movement
Species
Effect
Notes
Vol
Page
Ref
Vasotocin
Sleep (quiet state)
Rat
Increased
9
487
230
Vasotocin Vasotocin VIP
Sleep (REM) Sleep (REM) Motor act.
Human Human Rat
Increased Increased Increased
Neonates; dose, sleepstage interactions Narcoleptics Prepubertal boys ICV; enhanced by pargyline, suppressed by CCK-8
1 2 6
281 245 53
418 419 254
VIP VIP
Running Sleep
Rat Rat
Decreased Decreased
1 6
383 53
408 254
VIP VP VP
Sleep (REM) Exploratory behavior Exploratory behavior
Cat Rat Rat
Increased Decreased Increased
5 8 2
837 785 437
154 494 205
VP
Motor act.
Mouse
Increased
5
535
351
VP VP
Motor act. Motor act.
Rabbit Rat
No change Increased
6 15
2S 97 229
229 52
VP
Motor act.
Rat
Decreased
4
717
586
VP
Motor act.
Rat
Decreased
13
17
547
VP
Sleep (slow wave)
Human
Increased
17
1253
424
VP analog DGAVP Z-L-glutamyl-L-leucine
Exploratory behavior Jumping
Rat Mouse
Decreased Delayed
8 4
785 417
494 295
Z-L-glycyl-L-proline
Jumping
Mouse
Delayed
4
417
295
Z-L-leucyl-L-glycine
Jumping
Mouse
No change
4
417
295
Z-L-prolyl-L-leucine
Jumping
Mouse
No change
4
417
295
Z-L-prolyl-D-leucine
Jumping
Mouse
Delayed
4
417
295
ICV; pentobarbitalinduced sleep ICV Also increased latency to move and decreased postural freezing ICV; effect enhanced by co-admin. of amastatin Admin. to neonates, testing in adults In VP-deficient and controls Intrathecal admin.; blocked by V1 antagonist Elderly; subchronic treatment Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping
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Table 2 Addictive behavior Peptide
Behaviour
Species
Effect
ACTH fragment (1–24)
Self-admin. damphetamine Alcohol intake
Rat
Decreased
Rat
Decreased
Aminosenktide
Alcohol-aversive reactions
Rat
Decreased
Bombesin Bombesin
Alcohol intake Alcohol-induced righting impairment Apomorphineinduced yawning Alcohol intake
Rat Rat
Decreased Impaired
Rat
Aminosenktide
Casomorphin analogs CCK-8 CTOP Cyclo(Leu-Gly) Dermorphin analog TDAPA Dermorphin analog TDAPG-NH2 Endorphin () Enkephalin analog DAME Enkephalin (Leu) F8Fa F8Fa F8Fa F8Fa and analogs F8Fa antagonist daY8Ra F8Fa, antibody against FMRa and analogs FMRa and analogs FMRFa FMRFa analog F8Fa Galanin Litorin L-pyroglutamyl-L-leucine
Met-enkephalin-FMRFa chimeric peptide MIF-1
Vol
Page
Ref
5
659
307
19
1557
438
19
1557
438
ICV
6 2
2S 103 1S 99
302 187
Decreased
ICV
13
69
464
Rat
Decreased
14
1193
74
Cocaine-induced seizures Morphine-induced analgesia Morphine tolerance
Rat
Attenuated
Effect blocked by antagonist MK-329 ICV
18
1189
57
Rat
10
767
44
Mouse
Inhibited morphine tolerance No effect
11
139
78
Morphine tolerance
Mouse
No effect
11
139
78
Alcohol-induced righting impairment Ethanol preference and intake Ethanol preference and intake Morphine abstinence syndrome Morphine tolerance
Rat
Impaired
2
1S 99
187
Rat
Decreased
10
697
471
Rat
Decreased
10
697
471
Rat
No change
14
1271
459
Rat
No change
14
1271
459
Morphine withdrawal behavior Morphine abstinence syndrome Morphine abstinence syndrome Morphine abstinence syndrome Morphine abstinence syndrome Morphine abstinence syndrome Morphine withdrawal behavior Morphine withdrawal syndromes Opiate withdrawal behavior Alcohol intake Morphinewithdrawal-induced jumping Morphine tolerance
Rat
Precipitates
11
277
334
Rat
Decreased
12
1011
325
Rat
Decreased
ICV; morphinedependent rats ICV; in F8Fa-treated rats
12
1011
325
Rat
Decreased
11
969
335
Rat
Increased
17
83
484
Rat
Increased
14
731
337
Rat
Precipitates
14
47
338
Rat
Potentiated withdrawal
ICV; morphinedependent rats Third ventricle admin.; morphine-dependent rats Third ventricle admin.; morphine-dependent rats Third ventricle admin.; morphine-dependent rats ICV or IP
20
1211
539
Rat
No change
15
1303
237
Rat Mouse
Decreased Delayed
6 4
2S 103 417
302 295
Mouse
Attenuates tolerance
20
471
214
Rat
Inhibited tolerance
6
7
440
EtOH-induced sleep
Notes
ICV or admin to nucleus basalis magnocellularis; alcohol-preferring rats ICV or admin to nucleus basalis magnocellularis; alcohol-preferring rats; in alcohol-naive but not alcohol-experienced
ICV, intrathecal or SC; antinociceptive effect ICV, intrathecal or SC; antinociceptive effect ICV
Chronic ICV admin. with morphine Chronic ICV admin. with morphine ICV
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Table 2 (continued) Peptide
Behaviour
Species
Effect
MIF-1
Morphine-induced analgesia Morphine abstinence syndrome
Rat Rat
Inhibited morphine tolerance Increased
Alcohol-induced righting impairment Morphine tolerance Alcohol intake Apomorphineinduced yawning Apomorphineinduced yawning EtOH-induced sleep Alcohol intake
Rat
Impaired
Mouse Hamster Rat
Inhibited tolerance Increased Decreased
Rat
Decreased
Rat Rat
Inhibited tolerance Decreased
Alcohol-induced righting impairment Alcohol intake
Rat
Reversed
Rat
Decreased
Rat
Improved
Rat
Decreased
VP
Alcohol-induced righting impairment Apomorphineinduced fighting Alcohol intake
Rat
Decreased
VP (Lys)
Alcohol intake
Rat
Reversed
VP (Lys)
Alcohol pref.
Rat
Reversed
VP analog DGAVP
Alcohol intake
Rat
Reversed
VP analog DGAVP
Alcohol pref.
Rat
Reversed
VP analog DGAVP
Alcohol intake
Rat
Decreased
Z-L-glutamyl-L-leucine
Morphinewithdrawal-induced jumping Morphinewithdrawal-induced jumping Morphinewithdrawal-induced jumping Morphinewithdrawal-induced jumping Morphinewithdrawal-induced jumping
Mouse
Neuropeptide FF
Neurotensin NG-nitro-L-arginine NPY NT NT analog [D-Trp11]NT Oxytocin PG-KII Substance P Tachykinin aminosenkide
TRH TRH
Z-L-glycyl-L-proline
Z-L-leucyl-L-glycine
Z-L-prolyl-leucine
Z-prolyl-D-leucine
Notes
Vol
Page
Ref
10
767
44
17
615
336
2
1S 99
187
ICV
17 9 12
619 1389 755
611 300 398
ICV
12
755
398
6 18
7 825
440 437
1
103
262
19
897
416
2
1S 99
187
9
539
441
4
359
382
6
669
453
6
669
453
6
669
453
6
669
453
6
677
102
Delayed
4
417
295
Mouse
Delayed
4
417
295
Mouse
No change
4
417
295
Mouse
No change
4
417
295
Mouse
Delayed
4
417
295
ICV; antagonized by nitric oxide synthase inhibitors ICV
ICV; alcohol-preferring animals ICV; in haloperidol treated animals Selective effect in lateral hypothalamus and nucleus basalis magnocellularis but not other brain areas ICV
Only in AVP deficient rats AVP deficient with enhanced alcohol intake AVP deficient with enhanced alcohol intake AVP deficient with enhanced alcohol intake AVP deficient with enhanced alcohol intake In roman high avoidance and control rats
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the spinal cord contains many peptides and peptide receptors thought to influence pain, including calcitonin gene related peptide, somatostatin, neuropeptide Y, galanin, and substance P. Opioids also exert one of their main actions at this level by inhibiting release of substance P from primary nociceptive fibers. The spinothalamic and spinoreticular tracts are the best known pathways for transmitting pain from spinal cord to brain, although they are by no means the only course. Several descending pathways also allow the brain to influence the response to pain, and different anatomic locations within the brain influence the response to pain in ways not linked directly to the perception of the stimuli. This presents a plethora of locations at which a peptide can act to influence pain-related behavior within the brain. A listing of pain-related behaviors follows in Table 3. The majority of these studies investigated a simple response to a nociceptive stimuli in the presence of a particular peptide. Therefore rather than listing each paper with the behavior expected described (lifted tail, licked paw, etc.) we have included the protocol used to study the pain behavior and whether the peptide had an analgesic or anti-analgesic effect.
2.4. Ingestive behaviors Regulation of the ingestion of food and water is perhaps the behavior most critical to the survival of an animal. Proper regulation of such behavior depends on communication between brain, gut, and fat tissues, a communication that often makes use of peptides. The OB peptide leptin has received the most attention in recent years, but it should not be forgotten that leptin acts by regulating the expression of other neuropeptides in the brain [2]. These peptides are also active in regulating feeding in pathways that are independent of leptin. Other peripheral satiety systems such as that for CCK [374] are also critical. Drinking, like feeding, is highly regulated by peptides. Osmometric thirst is stimulated by osmoreceptors in the anterior basal forebrain and is influenced by VP. Volumetric thirst on the other hand is regulated by the actions of AT-II in the subfornical organ. Salt appetite also is largely mediated by the AT-II subfornical organ system, although other peptides like CRF and ACTH likewise play a role by stimulating the action of mineralocorticoids on the medial amygdala. Still other peptides appear to simultaneously regulate the intake of both water and food [374], or to regulate intake of particular nutrients [150]. Some nutrients blur the line between food and liquid intake. In particular, alcohol ingestion has often been studied in conjunction with investigations of food and/or water intake. For this reason we present those studies here (Table 4) as well as in the addictive behaviors section.
2.5. Social and sexual behaviors In no field is the idea that biochemicals including peptides, may influence or determine behavior more controversial than in the area of social and sexual behaviors. Although the influence on behavior in humans is often not known, the critical role of peptides in regulating these behaviors in animals is well established. Many of the most well known effects of peptides on sexual and social behaviors involve stimulation of other hormones. Lordosis for example, is said to be controlled by estrogen and progesterone, but estrogen and progesterone secretion in turn is controlled by peptides in the hypothalamus and pituitary gland. More direct effects of peptides on the brain also have been observed. For female sexual behavior the ventromedial nucleus of the hypothalamus plays a particularly important role in peptide receptivity whereas in males the medial preoptic area may be more important [73]. The anatomy of aggressive behavior often centers around the periaqueductal gray matter and the ventral tegmental area, although many studies of the effects of peptides have been done systemically. Social interaction, at least in rodents, may depend on smell, and thus many peptide effects are seen in olfactory areas. Emotional responsiveness provoked in the amygdala is known to be critical to social interactions, and thus the actions of peptides may influence social behaviors in this area. Memory and pain also tie in strongly with social behaviors, which, as mentioned in other sections, leaves quite a wide role for peptides. These behaviours are listed in Table 5. 2.6. Learning and memory behaviors Almost all behaviors are modified by experience. Some behavioral studies, however, are established mainly for the purpose of investigating the effects of experience itself on a particular behavior, in short, how well that behavior is learned. Since peptides act as neurotransmitters and neuromodulators in the brain, it is only to be expected that peptides play a critical role in learning behaviors [487]. The quest for peptides that when exogenously administered alter the ability to perform a learned task has also been the target of much research, since these substances may offer therapeutic hope for dementia [40]. The complexities of behavioral studies of learning can be impressive, teasing out different states of memory, effects on attention, reward, or other motivation. Although the purpose is to investigate internal states, results are gleaned by observation of a particular behavior. For this reason we have organized the learning and memory table by protocol used and, when appropriate, by the particular phase of that protocol in which the behavior was observed (Table 6).
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Table 3 Nociceptive behavior Peptide
Protocol
Species
Effect
Notes
Vol
Page
Ref
ACTH
Ear withdrawal
Rabbit
Anti-analgesic
7
1095
588
AT II
Tail flick
Rat
Analgesic
20
335
445
Calcitonin
Hot plate
Mouse
Analgesic
6
3S 277
200
Calcitonin
Hot plate
Mouse
Analgesic
6
3S 277
200
Calcitonin
Hot plate
Rat
Analgesic
6
3S 273
72
Calcitonin Casomorphin ()
Hot plate Tail flick, hot water, and tail compression test
Rat Rat
Analgesic Analgesic
7 10
315 539
213 585
Casomorphin analogs
Vocalization test
Rat
Analgesic
15
457
465
Casomorphin analogs CCK
Vocalization test Hot plate
Rat Mouse
Analgesic Analgesic
13 18
483 409
466 587
CCK
Tail flick and hot plate
Rat
Anti-analgesic
9
1295
21
CCK analog Boc[Phe(2,6diMe)7]-CCK-7 CCK analog Boc-[Phe(2Me)7]-CCK-7 CCK analog Boc-[Phe(4Me)7]-CCK-7 CCK analog Boc[Phe(pentaMe)7]-CCK-7 CCK analog Boc[Phe(triMe)7]-CCK-7 CCK analog Boc-CCK-7 CCK-8 CCK-8
Hot plate
Mouse
Analgesic
Endorphin ()-induced analgesia ICV; effect antagonized by naloxone ICV, IP, and IV, but no effect SC ICV, IP, and IV, but no effect SC; Ach-induced writhing Intrathecal admin.; antagonized by 6OHDA, haloperidol ICV ICV; greater effect in spontaneously hypertensive rats than controls Effects antagonized by naloxone ICV ICV; analgesic effect antagonized by naloxone but not CCK antagonists Morphine-induced analgesia; effect seen in tail flick but not hot plate ICV
19
301
333
Hot plate
Mouse
Analgesic
ICV
19
301
333
Hot plate
Mouse
Analgesic
ICV
19
301
333
Hot plate
Mouse
No change
ICV
19
301
333
Hot plate
Mouse
Analgesic
ICV
19
301
333
Hot plate Hot plate Hot plate
Mouse Mouse Rat
Analgesic Analgesic Anti-analgesic
19 19 6
301 301 237
333 333 253
CCK-8
Tail flick
Rat
Anti-analgesic
ICV ICV ICV; endorphin-induced analgesia; effect blocked by frontal cortex lesion ICV; morphine- or endorphin-, but not DPDPE-induced, analgesia
16
1229
528
Ceruletide Ceruletide Ceruletide
Mouse Human Human
Analgesic Analgesic Analgesic
2 3 3
2S 61 955 955
132 508 508
Ceruletide analogs Cyclo (Leu-Gly)
Hot plate Pain reporting Tolerance to electricity and thermal pain Hot Plate Tail flick
Mouse Rat
2 10
2S 65 767
609 44
Dalargin
Hot plate
Mouse
Analgesic Inhibited morphine tolerance Analgesic
19
777
482
D-Arg dipeptides
Tail flick
Mouse
Analgesic
6
35
475
Not blocked by naloxone Not blocked by naloxone
Morphine-induced analgesia Carried across the bloodbrain barrier by poly(butyl)cyanoacrylate nanoparticles ICV; mildly antagonized by naloxone
(continued on next page)
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Table 3 (continued) Peptide
Protocol
Species
Effect
Notes
Vol
Page
Ref
Deltakephalin
Tail flick, hot water, and tail compression test
Rat
Analgesic
ICV; greater effect in spontaneously hypertensive rats than controls
10
539
585
Deltorphin Deltorphin II Deltorphin II
Hot plate Tail flick Tail flick
Snail Mice Mouse
Analgesic Analgesic Analgesic
18 17 17
703 1415 619
544 45 611
Deltorphin II analog DADELT II Dermorphin
Tail flick
Rat
Analgesic
13
1123
251
Tail flick
Rat
Analgesic
6
3S 165
59
Tail flick Hot plate and tail flick Tail flick Flinch-jump after foot shock Pain reporting Pain reporting Tail flick Tail flick Hot plate Hot plate Ear withdrawal Hot plate
Rat Rat Mice Rat
Analgesic Analgesic Analgesic Analgesic
ICV; in dermorphin or morphine-tolerant-rats Intrathecal admin. ICV ICV ICV
13 3 17 3
1123 745 1415 41
251 492 45 426
Human Human Rat Rat Slug Slug Rabbit Rat
Analgesic Analgesic Analgesic Anti-analgesic No change No change Analgesic Analgesic
Intrathecal admin. Intrathecal admin. Intrathecal admin. ICV
8 8 6 6 10 10 7 6
191 191 1015 1015 9 9 1095 237
580 580 449 449 116 116 588 253
Hot plate and writhing tests Tail flick Hot plate Hot plate
Mouse
Analgesic
20
865
96
Rat Mouse Slug
Analgesic Analgesic Analgesic
ICV; CCK-8 antagonizes
16 3 10
1229 697 9
528 323 116
Hot plate
Rat
Analgesic
Attenuated by magnetic field
7
449
282
Enkephalin analog DAGO Enkephalin analog DALCE Enkephalin analog DAMA
Hot plate Hot plate Hot plate
Slug Rat Rat
Analgesic Analgesic Analgesic
10 10 1
9 319 139
116 71 257
Enkephalin analog DAMCK
Tail flick
Rat
Analgesic
20
1321
531
Enkephalin analog DPDPE
Hot plate
Snail
Analgesic
19
333
545
Enkephalin analog DPDPE Enkephalin analog DPDPE
Hot plate Tail flick
Snail Rat
Analgesic Analgesic
18 16
703 1229
544 528
Enkephalin analog metkephamide Enkephalin analog DAMGO Enkephalin analog DAMGO FMRFa
Tail flick
Rat
Analgesic
3
869
68
Tail flick
Rat
Analgesic
15
1567
608
Tail flick
Rat
Analgesic
20
1321
531
Hot plate
Rat
No change
10
735
64
Tail flick
Mouse
Analgesic
19
1171
444
Dermorphin Dermorphin analogs DPDPE Dynorphin Dynorphin frag (1–10) Dynorphin frag. (1–13) Dynorphin frag. (1–13) Dynorphin frag. (1–13) Dynorphin frag. (1–17) Dynorphin frag. (1–8) Endorphin () Endorphin () Endorphin () Endorphin () Enkephalin Enkephalin analog DADLE Enkephalin analog DAGO
FMRFa analog [D-Met2]-FMRFa
ICV ICV; not inhibited by NOS inhibitors Intrathecal admin.
ICV ICV; effects enhanced by frontal cortex ablation ICV
ICV ICV; antagonized by taurine and GABA Effect antagonized by naloxone pre-treatment, less so by post-treatment Some cross with pulsedmagnetic field-induced analgesia ICV; CCK-8 does not antagonize Antagonized by naloxone
Effect antagonized by naloxone Morphine-induced analgesia ICV; effect antagonized by antisense to the opioid mu receptor or by antisense to the Gi2␣ G-protein subunit
(continued on next page)
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Table 3 (continued) Peptide
Protocol
Species
Effect
Notes
Vol
Page
Ref
FMRFa analog A-18Fa
Hot plate
Mouse
Anti-analgesic
12
235
283
FMRFa analog DNS-RFa
Hot plate
Rat
Anti-analgesic
10
735
64
FMRFa analog F8Fa
Hot plate
Mouse
Anti-analgesic
12
235
283
FMRFa analog F8Fa
Hot plate
Mouse
Anti-analgesic
13
603
280
Hemorphin-4
Hot water; tail flick
Mouse
Analgesic
10
747
125
Hemorphin-5
Hot water; tail flick
Mouse
Analgesic
10
747
125
Kentsin
Hot plate and abdominal stretch
Rat
Analgesic
8
613
185
Lys (CH2NH)-Trp(Nps)OMe Lys-Trp
Tail flick (warm water)
Mouse
Analgesic
ICV; defeat-induced analgesia Morphine-induced analgesia ICV; defeat-induced analgesia ICV; morphine- or stress-induced analgesia ICV; naloxone antagonizes ICV; naloxone antagonizes With ICV and intrathecal admin. but not IV; antagonized by naloxone ICV; antagonized by naloxone
13
63
133
Response to heat and electricity Tail flick (warm water)
Mouse and rat
Analgesic
7
39
191
Mouse
Analgesic
13
63
133
20
471
214
Defeat-induced analgesia Defeat-induced analgesia Morphine-treated Rotation-induced analgesia Warm water stressinduced analgesia; no effect with cold waterinduced analgesia Morphine-induced analgesia
6 7 1 10
165 1007 293 493
542 279 147 322
6
1109
190
10
767
44
Morphine-induced analgesia ICV ICV ICis; blocked by GABA mimetics Morphine, but not deltorphin II, tolerance
3
433
121
10 7 8
539 1095 905
585 588 552
17
619
611
7 10 18 4
995 435 1223 517
49 167 473 249
18
1223
473
5 20
747 865
291 96
7 18 9
1095 93 619
588 53 92
3
851
519
Lys-Trp(Nps) Met-enkephalin-FMRFa chimeric peptide MIF-1 MIF-1 MIF-1 MIF-1
Tail flick
Mouse
Analgesic
Hot Hot Hot Hot
Mouse Mouse Mouse Vole
Anti-analgesic Anti-analgesic Anti-analgesic Anti-analgesic
MIF-1
Tail flick
Rat
Anti-analgesic
MIF-1
Tail flick
Rat
MIF-1
Tail flick
Rat
Inhibited morphine tolerance Delayed onset
Morphiceptin MSH Neo-Kyotorphin
Ear withdrawal Tail pinch
Rat Rabbit Mouse
Analgesic No change Analgesic
NG-nitro-L-arginine
Tail flick
Mouse
Inhibited tolerance
N-hexanol-5HTP-5HTP NT NT NT
Tail flick Hot plate Paw pressure test Tail flick
Rat Mouse Mouse Mouse
Analgesic Analgesic Analgesic Analgesic
NT analog Eisai hexapeptide OT Peptide E
Paw pressure test
Mouse
Analgesic
Rat Mouse
Analgesic Analgesic
Somatostatin Somatostatin analog CTAP SP
Tail flick Hot plate and writhing tests Ear withdrawal Tail flick (warm water) Ear withdrawal
Rabbit Mouse Rabbit
No change Analgesic Analgesic
SP
Hot plate
Mouse
Analgesic
plate plate Plate plate
ICV; antagonized by naloxone
ICV ICV Intrathecal admin.; variable effect ICV ICV ICV; only at very high doses ICV ICV; no effect of frag. (1–9) ICV
(continued on next page)
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Table 3 (continued) Peptide
Protocol
Species
Effect
Notes
Vol
Page
Ref
SP
Hot plate
Mouse
Analgesic
Naloxone antagonized effect; SP also antagonized immobilization but not foot shock analgesia
4
763
219
SP SP
Hot plate Motor response and vocalization after electroshock Shock threshold Motor response and vocalization after electroshock Hot plate
Mouse Rat
Analgesic Analgesic
ICV
4 7
31 419
218 390
Cat Rat
Analgesic Analgesic
Intrathecal admin. ICV
5 7
477 419
592 390
Mouse
Analgesic
Naloxone antagonized effect; no effect on stress-induced analgesia ICV
4
763
219
7
419
390
Immobilization but not foot shock analgesia Foot shock-induced analgesia ICV or IV; cold swiminduced analgesia
4
763
219
5
635
69
8
299
79
11
897
258
8
299
70
13
63
133
13
63
133
7 13
1007 1295
279 281
10
493
322
20
971
38
15 15
1508 1567
196 608
5
747
291
6
621
51
7
111
50
SP SP analog DiMe-C7
SP frag. (1–7)
SP frag. (4–11)
Rat
Analgesic
SP frag. (7–11)
Motor response and vocalization after electroshock Hot plate
Mouse
Anti-analgesic
TRH
Tail flick
Rat
Analgesic
TRH
Tail flick
Rat
Analgesic
TRH analog RX 77368
Rat
Analgesic
TRH analog RX 77368
Hot water tail immersion Tail flick
Rat
Analgesic
Trp(Nps)-Lys
Tail flick (warm water)
Mouse
Analgesic
Trp(Nps)(COCH2)(R,S)Lys Tyr-MIF-1 Tyr-MIF-1
Tail flick (warm water)
Mouse
Analgesic
Hot plate Hot plate
Mouse Mouse
Anti-analgesic Anti-analgesic
Tyr-MIF-1
Hot plate
Vole
Anti-analgesic
Tyr-W-MIF-1
Tail flick
Rat
Analgesic
Tyr-W-MIF-1 Tyr-W-MIF-1 and cyclic analogs VP
Tail flick Tail flick
Rat Rat
Analgesic Analgesic
Tail flick
Rat
Analgesic
VP
Tail flick
Rat
Analgesic
VP
Tail flick
Rat
Analgesic
VP VP
Tail flick Tail flick and flinchjump Tail flick
Rat Rat
Analgesic Analgesic
ICV but not SC
13 3
17 613
547 292
Rat
Analgesic
ICV
5
747
291
VP analog DDAVP
ICV; cold swim-induced analgesia ICV; antagonized by naloxone ICV; naloxone antagonizes Defeat-induced analgesia Morphine- or stressinduced analgesia Rotation-induced analgesia ICV; no tolerance, but cross-tolerance with morphine
ICV; not antagonized by naloxone ICV; enhanced by neonatal MSG ICV; paraventricular nucleus lesions block effect
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2197
Table 4 Ingestive behaviors Peptide
Behaviour
Species
Effect
[D-Ser , Leu ]enkephalin-Thr6 Acetorphan
Food intake
Rabbit
Decreased
Milk intake
Cat
Increased
ACTH (1–24)
Food intake
Rat
Decreased
Amylin
Food intake
Mouse
Decreased
Amylin Amylin
Food intake Food intake
Rat Rat
Decreased Decreased
Amylin Amylin
Food intake Food intake
Rat Rat
Decreased Decreased
Amylin
Food intake
Rat
Decreased
Amylin
Food intake
Rat
Decreased
Amylin
Food intake
Rat
Decreased
Amylin
Food intake
Rat
Decreased
Amylin
Saccharin preference
Rat
No change
Amylin Amylin ANF III Anorexigenic peptide Anorexigenic peptide Anorexigenic peptide AT antagonist sarthran AT II AT II
Water intake Water intake Water intake Food intake Food intake Water intake Water intake Salt intake Water intake
Rat Rat Sheep Dog Rat Rat Rat Rat Dog
Decreased No change Decreased Decreased Decreased Decreased Increased Increased Increased
AT AT AT AT AT AT
II II II II II II
Water Water Water Water Water Water
intake intake intake intake intake intake
Hamster Pigeon Rat Rat Rat Rat
Increased Increased Increased Increased Increased Increased
AT AT AT AT
II II II II
Water Water Water Water
intake intake intake intake
Rat Rat Rat Rat
Increased Decreased Increased Increased
Water intake
Rat
Increased
2
AT II
5
Notes
In sham feeding but in not non fasted ICV; also blocked stimulation by opiate agonists ICV or IP; in control, fooddeprived, diabetic, and insulin-treated animals Intra-hypothalamic admin. Intra-hypothalamic admin.; rat amylin more effective than human Vagotomized and controls Effect blocked by histamine antagonists, but not serotonin agonists or antagonists Effect blocked by amylin receptor antagonist In diabetic and control rats; effect antagonized by CGRP antagonist in nondiabetic animals ICV; not influenced by area postrema/nucleus of the solitary tract lesion Effect attenuated by lesion of area postrema/nucleus of solitary tract Intra-hypothalamic admin.; conditioned taste aversion Intra-hypothalamic admin. ICV Sham-feeding ICV infusion in females ICV infusion in females ICV ICV Alkalotic animals unresponsive ICV ICV Attenuated by vagotomy Effect blocked by ANF Greater in spontaneously hypertensive rats Sinoaortic-denervated Subfornical organ lesion Antagonized by ANF Synergistic effects with delta2 opioid agonists ICV; intraoral and free drinking measured
Vol
Page
Ref
7
745
208
10
967
17
7
834
435
12
865
370
13 12
961 919
79 18
16 17
457 1317
326 328
17
119
327
18
643
329
19
1533
330
19
309
331
13
961
79
13 19 12 2 4 4 10 3 9
961 1711 77 2S 235 85 85 261 493 1S 201
79 15 342 288 383 383 223 165 298
4 3 4 6 7 9
103 631 159 1S 145 373 979
358 130 491 212 388 595
10 10 11 18
103 423 837 241
266 265 393 606
19
171
241
(continued on next page)
2198
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 4 (continued) Peptide
Behaviour
Species
Effect
Notes
Vol
Page
Ref
AT II
Water intake
Rat
Increased
19
149
436
AT III
Water intake
Rat
Increased
9
979
595
AT IV
Water intake
Rat
No change
9
979
595
Atrial natriuretic factor Atriopeptin II Avian pancreatic polypeptide BBS BBS
Water intake Water intake Food intake
Rat Rat Chicken
Blocked AT effect Decreased Increased
ICV; effect antagonized by tachykinins Greater in spontaneously hypertensive rats In control and spontaneously hypertensive ICV; AT II-treated AT-stimulated drinking ICV
7 11 9
373 837 449
388 393 141
Alcohol intake Carbohydrate intake
Rat Rat
Decreased Decreased
6 11
2S 103 221
302 346
BBS BBS
Fat intake Feeding interval
Rat Rat
Decreased Decreased
11 12
221 761
346 182
BBS BBS BBS
Food intake Food intake Food intake
Baboon Dog Hamster
Decreased Decreased Decreased
4 5 7
687 675 1079
594 316 24
BBS
Food intake
Mouse
Decreased
9
221
490
BBS BBS BBS BBS BBS
Food Food Food Food Food
intake intake intake intake intake
Rat Rat Rat Rat Rat
Decreased Decreased Decreased Decreased Decreased
2 1 2 2 2
431 131 2S 179 261 409
491 339 198 314 498
BBS BBS
Food intake Food intake
Rat Rat
Decreased Decreased
3 3
61 61
581 581
BBS BBS
Food intake Food intake
Rat Rat
Decreased No change
3 5
553 41
199 589
BBS
Food intake
Rat
Decreased
5
547
39
BBS
Food intake
Rat
Decreased
5
481
94
BBS BBS
Food intake Food intake
Rat Rat
Decreased Decreased
5 6
607 1249
131 527
BBS
Food intake
Rat
Decreased
8
237
303
BBS
Food intake
Rat
Decreased
9
729
128
BBS
Food intake
Rat
Decreased
13
1215
99
BBS BBS
Food intake Food intake
Rat Rat
Decreased Decreased
16 17
903 119
523 327
BBS
Food intake
Rat
Decreased
17
119
327
BBS
Food intake
Rat
Decreased
18
1465
500
BBS
Food intake
Rat
Decreased
19
1407
101
Up to 2 hr post IP admin.; time course different for other macronutrients 4th ventricle admin.; blocked by antagonist In control and vagotomized In short- but not long-day photoperiod Greater effect in old than young
ICV or IP Insulin-induced feeding In controls and vagotomized Ventromedial hypothalamus lesions Admin. to hypothalamus, caudate putamen, olfactory tubercle In control and dorsomedial hypothalamic lesioned Sham-feeding, effect inhibited by proglumide ICV Effect blocked by GI neural disconnect In multiple brain sites; strongest in hypothalamus Nucleus of the tractus solitarius admin. ICV; animals developed tolerance after 8 days Effect partially blocked by amylin receptor antagonist Effect not blocked by amylin receptor antagonist Sham feeding and real feeding Weaker effect in high fat diet
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2199
Table 4 (continued) Peptide
Behaviour
Species
Effect
Notes
Vol
Page
Ref
BBS
Food intake
Rat
Decreased
19
309
331
BBS BBS
Food intake Food intake
Rat Rat
Increased Decreased
20 9
437 1S 233
572 260
BBS BBS BBS
Food intake Food intake Food seeking (bar press) Food seeking (bar press) Food seeking (bar press) Liquid food intake Milk intake
Turkey Wolf Rat
Decreased Decreased Decreased
Effect attenuated by lesion of area postrema/nucleus of solitary tract Amygdala admin. In IP or nucleus tractus solitarius, but not nucleus accumbens, admin. ICV and IV
10 7 12
275 969 1435
140 375 55
Rat
Decreased
13
343
347
Rat
Decreased
4
1
242
Rat Rat
Decreased Decreased
2 12
61 761
301 182
Mouse
Decreased
11
265
181
BBS
Milk seeking (bar press) Protein intake
Rat
Decreased
11
221
346
BBS BBS
Saccharin intake Salt intake
Rat Rat
No change Decreased
2 17
61 951
301 183
BBS BBS BBS BBS BBS BBS
Taste aversion Water intake Water intake Water intake Water intake Water intake
Rat Pigeon Pigeon Rat Rat Rat
No change Increased Increased No change Decreased Decreased
BBS BBS BBS BBS antagonist [D-Phe6, Leu13Cpal4]BBS(6–14) BBS antagonist [D-Phe6, Leu13Cpal4]BBS(6–14) BBS antagonist [D-Phe6, Leu13Cpal4]BBS(6–14) Calcitonin Calcitonin Calcitonin
Water intake Water intake Water intake Food intake
Rat Rat Turkey Rat
Food intake
BBS BBS BBS BBS BBS
Rate-dependent effect Capsaicin-treated and controls
4th ventricle admin.; blocked by antagonist
Up to 30 min post IP admin.; time course different for other macronutrients Sodium-deficient animals; in sham- and normaldrinking conditions
3 3 6 12 2 5
61 631 3S 181 1435 2S 179 41
581 130 129 55 198 589
Decreased Decreased Decreased No change
ICV ICV Rate dependent effect ICV or IP In hypothalamus but not caudate putamen, olfactory tubercle ICV ICV ICV and IV Neonates
5 6 10 14
607 3S 181 275 845
131 129 140 431
Rat
Increased
Only on postnatal day 15
17
1119
356
Water intake
Rat
No change
Neonates
14
845
431
Food intake Food intake Food intake
Dog Hamster Mouse
Decreased Decreased Decreased
In control and vagotomized
5 7 3
675 1079 17
316 24 373
Calcitonin Calcitonin
Food intake Food intake
Mouse Rat
Increased or decreased Decreased
9 7
221 729
490 577
Calcitonin Calcitonin
Food intake Food intake
Rat Rat
Decreased Decreased
3 5
749 861
551 297
Normal and genetically obese, diabetics Age dependent effect In control, lactating, and weaning SC or ICV injection
(continued on next page)
2200
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 4 (continued) Peptide
Behaviour
Species
Effect
Notes
Vol
Page
Ref
Calcitonin
Food intake
Rat
Decreased
9
729
128
Calcitonin
Food intake
Rat
Decreased
Nucleus of the tractus solitarius admin. Effect attenuated by lesion of area postrema/nucleus of solitary tract
19
309
331
Calcitonin Calcitonin Casomorphins Casomorphins
Food intake Water intake Carbohydrate intake Fat intake
Wolf Rat Rat Rat
No change Decreased Decreased Increased
7 3 19 19
969 749 325 325
375 551 319 319
CCK CCK
Carbohydrate intake Fat intake
Rat Rat
Decreased Decreased
11 11
221 221
346 346
CCK CCK CCK
Food intake Food intake Food intake
Dog Hamster Mouse
Decreased Decreased Decreased
5 4 9
675 103 221
316 358 490
CCK
Food intake
Mouse
Decreased
18
1275
341
CCK
Food intake
Rat
Decreased
2
409
498
CCK CCK CCK CCK
Food Food Food Food
intake intake intake intake
Rat Rat Rat Rat
Decreased Decreased Decreased Decreased
3 3 3 5
553 535 535 41
199 108 108 589
CCK
Food intake
Rat
Decreased
5
547
39
CCK
Food intake
Rat
No change
6
1S 133
104
CCK
Food intake
Rat
Decreased
6
927
106
CCK
Food intake
Rat
Decreased
4
15
243
CCK
Food intake
Rat
Decreased
7
587
455
CCK CCK
Food intake Food intake
Rat Rat
Decreased Decreased
8 9
763 601
604 503
CCK
Food intake
Rat
Improved
15
913
158
CCK
Food intake
Rat
Decreased
17
119
327
CCK
Food intake
Rat
Decreased
18
643
329
CCK
Food intake
Rat
Decreased
19
1407
101
CCK
Food intake
Rat
Decreased
19
309
331
CCK
Food intake
Rat
Decreased
1
47
344
ICV but not SC injection ICV or IP; antagonized by enterostatin Up to 6 hr post IP admin.; time course different for other macronutrients In control and vagotomized ICV Greater effect in old than young Synergistic effect with leptin In controls but not vagotomized ICV In hypothalamus but not caudate putamen, olfactory tubercle Effect reduced by dorsomedial hypothalamic lesioned Nucleus tractus solitarius injection Effect blocked by PVN lesions Two-day continuous infusion Effect attenuated by capsaicin Control and VP-deficient Effect blocked by capsaicin in fourth ventricle or vagus, but not spinal cord or pyloric region No effect of sex; effects potentiated by estradiol Effect partially blocked by amylin receptor antagonist In diabetic and control rats; effect antagonized by CGRP antagonist in nondiabetic Weaker effect in high fat diet Effect attenuated by lesion of area postrema/nucleus of solitary tract Hypothalamic or IP; norepinephrine-induced feeding
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2201
Table 4 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
CCK
Rat
Increased
959
264
Rat
Decreased
4
1
242
Mice
No change
2
39
525
CCK
Liquid food intake
Mice
Decreased
2
39
525
CCK
Number of meals
Rat
Decreased
4
15
243
CCK
Protein intake
Rat
Decreased
11
221
346
CCK
Sugar intake
Rat
Decreased
17
593
489
CCK
Time spent eating
Rat
Decreased
4
15
243
CCK
Water intake
Mice
Changed pattern
2
39
525
CCK
Water intake
Rat
Decreased
5
41
589
CCK CCK
Water intake Water intake
Rat Rat
No change No change
8 1
763 47
604 344
CCK agonist devazepide
Food intake
Rat
Increased
In amphetamine-treated but not controls Effect attenuated by food deprivation Normal and geneticallyobese mice Normal and geneticallyobese mice Two-day continuous infusion Up to 1 hr post IP admin.; time course different for other macronutrients Nucleus accumbens admin.; in animals with highbaseline sugar intake, but not in those with lowbaseline intake Two-day continuous infusion Genetically obese mice but not controls In hypothalamus but not caudate putamen or olfactory tubercle Control and VP-deficient Hypothalamic or IP; norepinephrine-induced consumption Effect during estrus, but not diestrus.
16
CCK
Food seeking (bar press) Food seeking (bar press) Inter meal interval
20
451
162
CCK agonist devazepide CCK analog Boc[Phe(2,6diMe)7]CCK-7] CCK analog Boc[Phe(2-Me)7]-CCK-7 CCK analog Boc[Phe(4-Me)7]-CCK-7 CCK analog Boc[Phe(pentaMe)7]CCK-7 CCK analog Boc[Phe(triMe)7]-CCK-7 CCK analog Boc-CCK-7 CCK analog ceruletide CCK analog JMV 170
Meal frequency Food intake
Rat Mouse
No change Decreased
20 19
451 301
162 333
Food intake
Mouse
Decreased
19
301
333
Food intake
Mouse
Decreased
19
301
333
Food intake
Mouse
Decreased
19
301
333
Food intake
Mouse
Decreased
19
301
333
Food intake Food intake Food intake
Mouse Human Rat
Decreased Decreased Decreased
19 3 11
301 607 873
333 506 410
CCK analog JMV 180
Food intake
Rat
Decreased
11
873
410
CCK antagonist L364,718 CCK antagonist L365,260 CCK antagonist L365,260
Food intake
Rat
Increased
40 min IV infusion After ICV but not IP admin. After ICV but not IP admin. CCK-A selective antagonist
12
1215
448
Food intake
Rat
No change
CCK-A selective antagonist
12
1215
448
Sugar intake
Rat
Decreased
IP and nucleus accumbens admin.; CCK-B receptor selective drug; high-sugar consuming rats
17
593
489
CCK
(continued on next page)
2202
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 4 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
CCK antagonist L365,260
Sugar intake
Rat
Increased
17
593
489
CCK frag. CCK-4
Rat
No change
19
849
376
Rat
Induced aversion
ICV, IV, or IP
19
849
376
Rat
Induced conditioning
16
539
Rat
Induced aversion
17
483
377
CCK-8
Taste aversion conditioning Taste aversion conditioning Taste aversion conditioning Taste aversion conditioning Alcohol intake
IP and nucleus accumbens admin.; CCK-B receptor selective drug; low-sugar consuming animals ICV, IV, or IP
Rat
Decreased
14
1193
74
CCK-8 CCK-8
Food intake Food intake
Chicken Dog
Decreased Decreased
3 8
739 651
142 468
CCK-8
Food intake
Hamster
Decreased
7
1079
24
CCK-8 CCK-8
Food intake Food intake
Human Mouse
Decreased
3 8
133 845
507 526
CCK-8 CCK-8 CCK-8 CCK-8
Food Food Food Food
Mouse Rat Rat Rat
Decreased Decreased Decreased Decreased
19 2 2 5
301 431 261 481
333 491 314 94
CCK-8
Food intake
Rat
Decreased
7
729
577
CCK-8
Food intake
Rat
Decreased
7
729
577
CCK-8
Food intake
Rat
Decreased
8
223
495
CCK-8 CCK-8
Food intake Food intake
Rat Rat
Decreased Decreased
9 9
1s 207 1093
481 252
CCK-8 CCK-8
Food intake Food intake
Rat Rat
Decreased Decreased
9 14
989 1193
460 74
CCK-8
Food intake
Rat
Decreased
17
1313
574
CCK-8
Food intake
Rat
Decreased
19
977
89
CCK-8
Food intake
Rat
Decreased
5
97
560
CCK-8 CCK-8
Food intake Food intake
Sheep Sheep
Decreased Decreased
1 3
51 963
137 138
CCK-8 CCK-8
Food intake Food seeking (bar press)
Wolf Rat
Decreased Decreased
7 4
969 351
375 590
CCK-8
Milk seeking (bar press)
Mouse
Decreased
11
265
181
CCK frag. CCK-8 CCK frag. CCK-8 CCK frag. CCK-8
intake intake intake intake
Effect blocked by antagonist MK-329 ICV In ICV but not IV; fooddeprived and insulininduced feeding Greater effect in short-day photoperiod 30 min IV infusion Linear dose relationship in males but not females; greater effect in lean than in obese
Insulin-induced feeding Sham-feeding, effect inhibited by proglumide Greater in lactating mothers than after weaning Resistance to effect during lactation and weaning In control and tumorbearing Reversed by proglumide ICV; proglumide antagonized; no effect from atropine or naloxone NPY treated Effect blocked by antagonist MK-329 In free feeding old and young, but not fixed feeding young. Stronger effect with progesterone and estradiol Tumor-bearing and control animals 6 hr ICV infusion ICV; no effect in cisterna magna Effect altered by GABAergic, dopaminergic and opioid drugs
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2203
Table 4 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
CCK-8 CCK-8
Water intake Water intake
Chicken Dog
Decreased Decreased
3 8
739 651
142 468
CCK-8
Water intake
Mouse
Decreased
8
845
526
CCK-8 CCK-8S Ceruletide
Water intake Food intake Food intake
Mouse Rat Human
Increased Decreased No change
ICV In ICV but not IV; fooddeprived and insulininduced feeding In obese, but not lean, males In lean and obese females
8 17 8
845 355 455
526 576 77
Ceruletide
Food intake
Rat
Decreased
5
1149
499
CGRP
Food intake
Mouse
Decreased
17
511
369
CGRP CGRP
Food intake Food intake
Rat Rat
Decreased Decreased
5 19
861 1533
297 330
CGRP
Mouse
Decreased
17
511
369
CGRP CRF
Milk seeking (bar press) Water intake Food intake
Mouse Rat
No change Decreased
17 4
511 807
369 209
CRF CRF
Food intake Food intake
Rat Rat
Decreased Decreased
6 13
3S 53 879
392 226
CRF
Food intake
Rat
Decreased
20
509
571
CRF
Food seeking (bar press) Food intake
Monkey
Increased
In control and vagotomized, but not adrenalectomized ICV admin. ICV; tail-pinch-induced feeding ICV; not influenced by melanocortin antagonist ICV
10
1199
384
Goldfish
Increased
Serotonin-induced anorexia
19
505
136
Food intake
Rat
ICV; tail-pinch- or NPYinduced feeding
13
879
226
Ethanol preference and intake Food intake Food intake
Rat
Increase at low doses; decrease at high doses Decreased
10
697
471
Goldfish Rat
Increased Increased
17 7
421 711
135 332
Water intake Food seeking (bar press) Water intake Water intake Water intake Food intake Food intake Food intake
Rat Monkey
Increased Increased
18 10
241 1199
606 384
Pigeon Rat Rat Chicken Rat Rat
No change Decreased Increased No change Increased Increased
6 6 18 8 4 7
3S 181 3S 181 241 585 797 711
129 129 606 518 372 332
Food Food Food Food Food
Rabbit Rat Rat Rat Rat and mouse Rat Rat Rat
Increased Increased Increased Increased Increased
7 4 4 4 1
745 797 797 797 341
208 372 372 372 578
4 4 4
797 797 797
372 372 372
CRF antagonist ␣-helical CRF (9–41) CRF antagonist ␣-helical CRF (9–41) DAME DAMGO DADLE Deltorphin II Dermorphin Dermorphin Dermorphin DPDPE Dynorphin Dynorphin Dynorphin Dynorphin Dynorphin Dynorphin Dynorphin Dynorphin
A frag. frag. frag. frag.
(1–10) (1–11) (1–13) (1–13)
Dynorphin frag. (1–8) Dynorphin frag. (1–9) Dynorphin frag. (3–13)
intake intake intake intake intake
Food intake Food intake Food intake
No change No change Increased
Obese women on restricted diet With IP, but not ICV, admin. Not influenced by CCK receptor antagonist ICV; not influenced by area postrema/nucleus of the solitary tract lesion
Admin. to nucleus accumbens septi Interactions with AT II ICV; blocked by naloxone ICV ICV Interactions with AT II ICV; 2 day old ICV Admin. to nucleus accumbens septi
ICV; not reversible by naloxone
(continued on next page)
2204
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 4 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
EGF Eledosin Endorphin () Endorphin () Endorphin ()
Water intake Water intake Food intake Food intake Food intake
Sheep Pigeon Chicken Goldfish Pigeon
Decreased Increased Increased Increased Increased
9 3 9 17 5
1221 631 709 421 691
417 130 345 135 145
Endorphin ()
Food intake
Rat
Increased
3
421
313
Endorphin ()
Food intake
Rat
Increased
7
711
332
Endorphin ()
Water intake
Chicken
9
709
345
Endorphin () Enkephalis analog DAME Enkephalis analog DAME Enkephalis analog DAME Enkephalin (Leu)
Water intake Milk intake Sugar intake Water intake Ethanol preference and intake Sugar intake Water intake Food intake
Pigeon Cat Rat Rat Rat
Decrease at early times, followed by increase at late times No change Decreased No change Decreased Decreased
ICV ICV ICV Mu-receptor effect Pre-fed, ICV injections, only for avian peptide Hypothalamic PVN injection Admin. to nucleus accumbens septi ICV
5 10 10 10 10
691 967 697 697 697
145 17 471 471 471
Rat Rat Rat
No change Decreased Increased
10 10 4
697 697 287
471 471 349
Water intake
Rat
Increased
4
287
349
Food intake
Goldfish
Increased
17
421
135
Carbohydrate intake Fat intake
Rat Rat
No change Decreased
18 18
785 785
409 409
Enterostatin Enterostatin
Fat intake Fat intake
Rat Rat
Decreased Decreased
18 19
657 325
318 319
Enterostatin Enterostatin
Feeding rate Food intake
Rat Rat
Decreased Increased
19 17
557 885
321 451
Enterostatin
Food intake
Rat
18
1341
320
F8Fa
Food intake
Rat
Decreased at low doses but not high doses Decreased
17
353
379
FMRFa
Food intake
Rat
Decreased
6
847
278
Galanin Galanin Galanin
Carbohydrate intake Fat intake Fat preference
Rat Rat Rat
No change No change Increased
18 18 9
207 207 309
496 496 541
Galanin
Food intake
Rat
Decreased
11
995
304
Galanin Galanin Galanin
Food intake Food intake Food intake
Rat Rat Rat
Increased Increased Increased
15 15 17
1303 1267 237
237 497 25
Galanin
Food intake
Rat
Increased
9
309
541
Gastrin
Food intake
Rat
No change
5
41
589
Enkephalin (Leu) Enkephalin (Leu) Enkephalin analog DALA Enkephalin analog DALA Enkephalin analog DPEN Enterostatin Enterostatin
Pre-fed, ICV injection
PVN injection PVN injection; only when food available
ICV ICV; effect blocked by kappa opioid antagonist ICV ICV; antagonized by casomorphins PVN and amygdala admin. ICV; high fat foods; free fed rats Effect when admin. to PVN or amygdala but not other brain areas ICV; no additive effect with naloxone ICV; kappa opiate agonistinduced PVN admin. PVN admin. Paraventricular nucleus admin. After admin. to PVN and periventricular region, but not other hypothalamic areas ICV ICV; effects not sustained Effect blocked by mu antagonists in animals fed high-fat diet Paraventricular nucleus admin. In hypothalamus, caudate putamen, olfactory tubercle
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2205
Table 4 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
Gastrin Gastrin
Water intake Water intake
Rat Rat
Decreased Decreased
Hypothalamic admin. Admin. to caudate putamen, olfactory tubercle
5 5
41 41
589 589
Gastrin (1–17) Gastrin (1–17) Gastrin-related peptide
Food intake Water intake Milk seeking (bar press) Feeding interval Food intake Food intake Food intake Food intake
Rat Rat Mouse
Decreased Decreased Decreased
1 1 11
383 383 265
408 408 181
Rat Rat Rat Rat Rat
Increased Decreased Decreased Decreased Decreased
19 2 3 16 17
1439 2S 179 833 903 107
462 198 517 523 524
Gastrin-releasing peptide
Food intake
Rat
Decreased
18
1465
500
Gastrin-releasing peptide Gastrin-releasing peptide
Food intake Food intake
Rat Rat
No change Decreased
19 20
1439 731
462 16
Gastrin-releasing peptide
Milk intake
Rat
Decreased
20
737
463
Gastrin-releasing peptide
Salt intake
Rat
Decreased
17
951
183
Gastrin-releasing peptide
Sugar intake
Rat
Decreased
20
737
463
Gastrin-releasing peptide GHRH
Water intake Carbohydrate intake
Rat Rat
No change No change
3 15
833 1343
517 150
GHRH
Food intake
Rat
Increased
15
1343
150
GHRH
Food intake
Rat
Increased
16
595
558
GHRH
Food intake
Rat
Decreased
2
261
314
GHRH
Food intake
Rat
9
1S 35
557
GHRH
Protein intake
Rat
Increase at low doses; decrease at high doses Increased
15
1343
150
GHRH, antibody to
Carbohydrate intake
Rat
No change
16
7
149
GHRH, antibody to
Protein intake
Rat
Decreased
16
7
149
Glucagon
Food intake
Dog
Decreased
5
675
316
Glucagon Glucagon
Food intake Food intake
Hamster Mouse
No change Decreased
7 9
1079 221
24 490
Glucagon Glucagon Glucagon
Food intake Food intake Food intake
Rat Rat Rat
Decreased Decreased Increased
3 3 8
163 163 943
195 195 194
Gastrin-releasing Gastrin-releasing Gastrin-releasing Gastrin-releasing Gastrin-releasing
peptide peptide peptide peptide peptide
Synergistic effect with neuromedin B In real feeding but not sham feeding Sham feeding; Synergistic effect with pre-feeding Effect on free feeding but not intraoral intake Sodium-deficient animals; in sham- and normaldrinking Effect on free feeding but not intraoral intake Suprachiasmatic nucleus/ medial preoptic area admin. Suprachiasmatic nucleus/ medial preoptic area admin.; effect blocked by methylnaltrexone Suprachiasmatic nucleus admin.; light-cycle dependent effect Hypothalamic admin.; insulin-induced feeding ICV Suprachiasmatic nucleus/ medial preoptic areas admin.; effect blocked by methyl-naltrexone Suprachiasmatic/preoptic area admin.; feeding studied at dark onset Suprachiasmatic/preoptic area admin.; feeding studied at dark onset In control, but not vagotomized Greater effect in old than young Sham-feeding
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2206
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 4 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
Glucagon
Food intake
Rat
Decreased
Effect blocked by amylin receptor antagonist when given at dark onset
17
119
327
Glucagon frag. (1–21) Glucagon-like peptide-1(7–37) amide Insulin Insulin Insulin
Food intake Food intake
Rat Rat
No change Decreased
ICV; in lean and obese
8 19
943 869
194 248
Food intake Food intake Food intake
Rat Rat Rat
Streptozocin-diabetic rats
4 12 13
79 425 721
124 98 348
Insulin Interleukin-1
Food intake Food intake
Rat Rat
Increased Increased Decreased (returned to same as controls) Decreased Decreased
19 14
1549 175
156 553
Interleukin-1
Food intake
Rat
Decreased
17
517
502
Kassinin
Water intake
Rat
Decreased
10
125
422
Leptin Leptin
Food intake Food intake
Mouse Mouse
Decreased Decreased
20 18
595 1275
368 341
Leptin Litorin Litorin Litorin Litorin Litorin Litorin MIF-1
Food intake Alcohol intake Food intake Food intake Water intake Water intake Water intake Food intake
Rat Rat Rat Rat Pigeon Rat Rat Rat
Decreased Decreased Decreased Decreased Increased Decreased Decreased No change
19 6 2 5 6 6 5 15
1549 2S 103 2S 179 607 3S 181 3S 181 607 243
156 302 198 131 129 129 131 216
MIF-1 Motilin
Water intake Food intake
Rat Mouse
Decreased Increased
1 19
353 987
407 13
Motilin Motilin Motilin MSH (␣)
Food intake Food intake Water intake Food intake
Rat Rat Rat Rat
Decreased Increased Decreased Decreased
1 6 1 7
383 41 383 834
408 193 408 435
MSH frag. analog Lys-D-Pro-Thr MSH frag. Lys-Pro MSH frag. Lys-Pro-Val
Food intake
Rat
Decreased
14
175
553
Food intake Food intake
Rat Rat
Increased Increased
14 14
175 175
553 553
Neoendorphin (␣)
Food intake
Rat
Increased
7
711
332
Neurokinin A
Water intake
Rat
Decreased
11
339
423
Neuromedin B
Food intake
Rat
Decreased
17
107
524
Neuromedin B
Salt intake
Rat
No change
17
951
183
NPY NPY
Alcohol intake Amino acid deficient diet intake
Hamster Rat
Increased Decreased
9 19
1389 527
300 111
Hypoglycemic rats Antagonized by MSH frag. Lys-Pro-Val and frag. analog Lys-D-Pro-Thr ICV; effect blocked by NPY Neonates; AT II-induced thirst Genetically obese mice Synergistic effect with CCK Hypoglycemic rats
ICV ICV ICV ICV Peptide YY-induced hyperphagia ICV; attenuated by motilin receptor antagonist Fasted subjects ICV; also blocked stimulation by opiate agonists In IL-1-induced anorexia but not controls In IL-1-induced anorexia In IL-1-induced anorexia but not controls Admin. to nucleus accumbens septi Neonates; in control and AT-treated Synergistic effect with gastrin releasing peptide Sodium-deficient animals; in sham- and normaldrinking Anterior piriform complex admin.; threonineimbalanced model
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R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2207
Table 4 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
NPY NPY NPY
Carbohydrate intake Carbohydrate intake Carbohydrate intake
Rat Rat Rat
Increased Increased Increased
6 12 18
1205 1251 207
510 312 496
NPY NPY NPY
Fat intake Fat intake Food intake
Rat Rat Chicken
No change No change Increased
12 18 8
1251 207 585
312 496 518
NPY NPY
Food intake Food intake
Chicken Dog
Increased Decreased
Admin. to PVN PVN admin. PVN admin.; in carbohydrate-preferring, but not control, rats PVN admin. PVN admin. ICV; 2 day old; in animals that did not have convulsions ICV ICV; in insulin-treated, but not fasted or satiated
8 8
823 651
299 468
NPY NPY NPY
Food intake Food intake Food intake
Hamster Mouse Rat
Increased Increased Increased
9 10 5
1389 963 1025
300 177 315
NPY NPY
Food intake Food intake
Rat Rat
Increased Increased
7 9
1189 723
512 269
NPY
Food intake
Rat
Increased
9
1269
467
NPY
Food intake
Rat
Increased
9
989
460
NPY
Food intake
Rat
Increased
10
1283
81
NPY
Food intake
Rat
Increased
12
609
411
NPY NPY NPY
Food intake Food intake Food intake
Rat Rat Rat
Increased Increased Increased
12 13 14
1251 581 475
312 513 514
NPY NPY
Food intake Food intake
Rat Rat
Increased Increased
15 16
799 599
54 112
NPY
Food intake
Rat
Decreased
17
797
80
NPY
Food intake
Rat
Increased
17
797
80
NPY
Food intake
Rat
Increased
17
797
80
NPY
Food intake
Rat
Increased
17
517
502
NPY
Latency to onset of feeding Milk seeking (bar press) Protein and fat intake Protein intake Sugar intake Sugar intake Water intake
Rat
Decreased
9
1269
467
Mouse
Increased
12
1329
178
Rat Rat Rat Rat Chicken
No change Decreased Decreased Increased No change
6 12 20 20 8
1205 1251 601 601 823
510 312 543 543 299
NPY NPY NPY NPY NPY NPY
ICV Effect inhibited by naloxone and haloperidol PVN of hypothalamus ICV; satiated females; attenuated by adrenalectomy; no effect from CCK ICV; effect attenuated by hypothalamic transection ICV; attenuated by intravenous infusions of glucose, but not fructose Paraventricular hypothalamic admin.; effect blocked by pertussis toxin Sustained admin. to PVN of hypothalamus PVN admin. Hypothalamic admin. Hypothalamic perifornical admin.; similar effects at different times of day ICV Medial PVN and lateral perifornical hypothalamus admin. Hypothalamic admin.; tumor-bearing Hypothalamic admin.; constant infusion; tumorbearing Daily increasing doses; effect in control but not tumor-bearing ICV; in IL-1-treated and controls ICV; effect attenuated by hypothalamic transection ICV; also overcame taste aversion Admin. to PVN PVN admin. ICV; continuous infusion ICV; single admin. ICV
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2208
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 4 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
NPY
Water intake
Dog
Decreased
ICV; in insulin-treated, but not fasted or satiated
8
651
468
NPY NPY
Water intake Water intake
Hamster Rat
Increased Increased
9 5
1389 1025
300 315
NPY analog [Leu31, Pro34]NPY NPY analog [Leu31, Pro34]NPY NPY analog [Leu31, Pro34]NPY NPY analog [Leu31, Pro34]NPY NPY analog C2-NPY
Carbohydrate intake
Rat
Increased
12
1251
312
Fat intake
Rat
No change
12
1251
312
Food intake
Rat
Increased
12
1251
312
Protein intake
Rat
Decreased
12
1251
312
Food intake
Rat
13
581
513
NPY analog [Pro34]NPY NPY antagonist BW1229U91 NPY antibodies NPY antibody
Food intake Food intake
Rat Rat
Increased (slight effect) Increased No change
Analog is Y1 selective ICV; control and obese
13 20
581 367
513 583
Food intake Food intake
Rat Rat
Decreased Decreased
13 15
581 607
513 579
NPY frag. (1–30) NPY frag. (13–36)
Food intake Carbohydrate intake
Rat Rat
No change Decreased
15 12
799 1251
54 312
NPY frag. (13–36)
Food intake
Rat
Decreased
12
1251
312
NPY frag. (13–36)
Protein intake
Rat
Decreased
12
1251
312
NPY NPY NPY NPY NPY NPY NPY NPY NT
Food intake Carbohydrate intake Fat intake Food intake Food intake Protein intake Food intake Food intake Food intake
Mouse Rat Rat Rat Rat Rat Mouse Rat Rat
No change Increased No change Increased Increased Increased No change No change Decreased
10 12 12 12 13 12 10 13 4
963 1251 1251 1251 581 1251 963 581 493
177 312 312 312 513 312 177 513 511
NT
Food intake
Rat
Decreased
9
729
128
NT
Water intake
Rat
No change
4
493
511
OT
Food intake
Rat
Decreased
10
89
12
OT
Food intake
Rat
Decreased
12
113
405
OT OT
Food intake Food intake
Rat Rat
Decreased Decreased
17 17
1119 1119
356 356
OT
Latency to first meal
Rat
Increased
ICV In ventromedial hypothalamus and ventromedial and ventrolateral thalamus, but not other hypothalamic areas ICV PVN admin.; frag. is Y2 selective PVN admin.; frag. is Y2 selective PVN admin.; frag. is Y2 selective ICV PVN admin. PVN admin. PVN admin. Hypothalamic admin. PVN admin. ICV Hypothalamic admin. Hypothalamic PVN injection Nucleus of the tractus solitarius admin. Hypothalamic PVN injection IP or ICV; blocked by pretreatment with d(CH2)5Tyr(Me)-Orn8vasotocin Antagonist blocked effect, but had no effect on its own In neonates and adults In neonates and adults; effect greater in adults IP or ICV; blocked by pretreatment with d(CH2)5Tyr(Me)-Orn8vasotocin
10
89
12
frag. (20–36) frag. (2–36) frag. (2–36) frag. (2–36) frag. (2–36) frag. (2–36) frag. (26–36) free acid
Effect inhibited by naloxone and haloperidol PVN admin.; analog is Y1 selective PVN admin.; analog is Y1 selective PVN admin.; analog is Y1 selective PVN admin.; analog is Y1 selective Analog is Y2 selective
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2209
Table 4 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
OT
Time spent eating
Rat
Decreased
10
89
12
OT antagonist vasotocin PACAP PACAP
Food intake Food intake Food intake
Rat Mouse Rat
Increased Increased Decreased
17 13 16
1119 1133 1511
356 371 82
PACAP
Water intake
Rat
Decreased
16
1511
82
Pancreatic polypeptide Pancreatic polypeptide Pancreatic polypeptide
Food intake Food intake Food intake
Chicken Dog Human
Increased No change Decreased
IP or ICV; blocked by pretreatment with d(CH2)5Tyr(Me)-Orn8vasotocin Only on postnatal day 10 ICV Hypothalamic admin.; in NPY-treated and controls Hypothalamic admin.; in NPY-treated and controls Intracerebral inj.; 2 day old
8 5 14
585 675 497
518 316 42
Pancreatic polypeptide Pancreatic polypeptide Peptide YY Peptide YY Peptide YY
Food intake Food intake Carbohydrate intake Food intake Food intake
Rat Rat Rat Chicken Rat
Increased Decreased Increased Increased Increased
11 20 6 8 15
673 1445 1205 823 243
19 14 510 299 216
Peptide YY Peptide YY PG-KII
Protein and fat intake Water intake Alcohol intake
Rat Chicken Rat
No change No change Decreased
6 8 18
1205 823 825
510 299 437
PG-KII
Food intake
Rat
No change
18
825
437
PG-KII
Water intake
Rat
Decreased
18
825
437
Ranatensin Ranatensin Ranatensin Ranatesin Sauvagine
Water intake Water intake Water intake Food intake Food intake
Pigeon Rat Rat Rat Rat
Increased Decreased Decreased Decreased Decreased
6 6 5 5 6
3S 181 3S 181 607 607 3S 53
129 129 131 131 392
Somatostatin
Amino acid deficient diet intake
Rat
Biphasic time and dose-related-effect
19
527
111
Somatostatin Somatostatin Somatostatin Somatostatin analog SMS 201–995 Somatostatin antibody Tachykinin agonist NP-␥
Food Food Food Food
Dog Rat Rat Rat
No change Decreased Increased Increased
5 19 19 9
675 991 991 1S 211
316 477 477 117
Food intake Water intake
Rat Rat
Decreased Decreased
9 19
1S 211 149
117 436
Tachykinin aminosenkide
Alcohol intake
Rat
Decreased
19
897
416
Tachykinin aminosenkide
Food intake
Rat
Decreased
19
897
416
Tachykinin aminosenktide
Water intake
Rat
Decreased
19
897
416
intake intake intake intake
Prader-Willi syndrome; effect only seen in females Hypothalamic admin. ICV Admin. to PVN ICV ICV; antagonized by naloxone Admin. to PVN ICV ICV; alcohol-preferring animals ICV; alcohol-preferring animals ICV; alcohol-preferring animals; in AT-stimulated drinking, but not control, animals ICV ICV ICV ICV ICV, and SC in vagotomized animals Anterior piriform complex admin.; Threonineimbalanced model In dark phase In light phase ICV ICV ICV; AT-stimulated drinking Selective effect in lateral hypothalamus and nucleus basalis magnocellularis but not other brain areas Effect with admin. to stria terminalis but not other brain areas Effect with admin. to stria terminalis but not other brain areas
(continued on next page)
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Table 4 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
Tachykinin analog Suc-[Asp6,MePhe8]Substance P(6–11) Tachykinin analog Suc-[Asp6,MePhe8]Substance P(6–11) Tachykinins Tachykinins TRH Tuftsin
Food intake
Rat
Decreased
Alcohol-preferring animals
16
533
88
Food intake
Rat
Alcohol-preferring animals
16
533
88
Water intake Water intake Food intake Food seeking (bar press) Food intake
Pigeon Rat Dog Monkey
Decreased at low doses but not high doses Increased Decreased Decreased Increased
ICV ICV Sham-feeding ICV
6 6 2 10
3S 181 3S 181 2S 235 1199
129 129 288 384
Rabbit
Decreased
7
745
208
Food intake
Rat
Increased
20
1425
114
Tyr-MIF-1
Food intake
Rat
No change
15
243
216
Urotensin VIP VIP VP VP
Food intake Food intake Water intake Alcohol intake Water intake
Rat Rat Rat Rat Rat
6 1 1 4 12
3S 53 383 383 359 285
392 408 408 382 584
VP
Water intake
Rat
Decreased No change No change Decreased Decreased (returned to same as controls) Decreased
1
11
190
VP VP (Lys)
Water intake Alcohol intake
Sheep Rat
Decreased Reversed
9 6
1221 669
417 453
VP (Lys)
Alcohol preference
Rat
Reversed
6
669
453
VP analog DGAVP
Alcohol intake
Rat
Decreased
6
677
102
VP antagonists VP frag. DGAVP
Water intake Alcohol intake
Dog Rat
Decreased Reversed
8 6
1003 669
532 453
VP frag. DGAVP
Alcohol preference
Rat
Reversed
6
669
453
Tyr-D-Ala-Gly-(Me) Phe-Gly-ol Tyr-MIF-1
Neonates; stress-induced feeding Peptide YY-induced hyperphagia ICV admin.
Only in VP-deficient rats Food restricted; VPdeficient Hypothalamic graft of VPproducing cells in VPdeficient rats ICV VP-deficient with enhanced alcohol intake VP-deficient with enhanced alcohol intake In Roman high avoidance and control rats ICV VP-deficient with enhanced alcohol intake VP-deficient with enhanced alcohol intake
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2211
Table 5 Social and sexual behavior Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
A18Fa octa-decapeptide
Defeat behavior
Mouse
Increased
12
235
283
A18Fa, antibody to
Defeat behavior
Mouse
Decreased
12
235
283
ACTH
Sexual behavior
Newt
4
729
361
ACTH ACTH frag. (4–10) ACTH/MSH 4–10 analog ORG 2766 ACTH/MSH 4–10 analog ORG 2766 Amylin
Sexual behavior Sexual behavior Social interaction
Rat Newt Rat
Increased ICV; decreased IP Decreased Increased Increased
ICV; subordinate mice; possible anti-analgesic effect ICV; subordinate mice; possible analgesic effect Opposite effects ICV or IP
14 4 2
379 729 255
8 361 174
Social interaction
Rat
Increased
2
255
174
Sexual behavior
Rat
Decreased
20
379
90
CCK-4
Defensive behavior
Rat
Increased
20
383
575
CGRP
Sexual behavior
Rat
Increased
20
379
90
CRF CRF CRF Dynorphin Endorphin ()
Aggressive behavior Defensive behavior Social interaction Lordosis Lordosis
Mouse Mouse Mouse Rat Rat
8 8 8 8 8
935 935 935 309 309
353 353 353 427 427
Enkephalin (Met)
Aggressive behavior
Goldfish
Decreased Increased Decreased No change Dose-dependent increase or decrease Decreased
1
73
474
Enkephalin (Met)
Aggressive behavior
Fish
6
139
86
Enkephalin analog [D-Ala2-Met5]enkephalinamide
Defensive behavior
Cat
9
999
486
F8Fa
Defeat behavior
Mouse
Increased
12
235
283
F8Fa, antibody to
Defeat behavior
Mouse
Decreased
12
235
283
LHRH
Sexual behavior
Human
No change
2
1S 115
163
LHRH antagonist MCH
Sexual behavior Sexual behavior
Rat Rat
Decreased Increased
2 17
393 171
157 207
MIF-1 MIF-1 Morphiceptin
Aggressive behavior Aggressive behavior Lordosis
Mouse Mouse Rat
ICV
6 7 8
165 1007 309
542 279 427
MSH
Lordosis
Rat
ICV or SC
7
581
446
MSH (␣)
Aggressive behavior
Mouse
Decreased Decreased Dose-dependent inhibition or facilitation Increased or decreased Increased
1
69
400
MSH (␣)
Aggressive behavior
Rat
Increased
17
171
207
MSH (␣)
Sexual behavior
Newt
Increased
Aggression seen in mouse paired with MSHtreated animal Admin. to ventromedial nucleus; antagonized by MCH ICV
4
729
361
Increased at low dose, decreased at high dose Increased at low dose, decreased at high dose
ICV In familiar, but not in unfamiliar, surroundings
ICV; effect blocked by CGRP Ultrasound-induced defense behavior ICV; in rats with amylinsuppressed sexual behavior ICV ICV ICV ICV ICV ICV; interaction with melatonin
Admin. to midbrain periaqueductal gray; electrical stimulationelicited defense behavior ICV; subordinate mice; possible anti-analgesic effect ICV; subordinate mice; possible analgesic effect Married men with secondary impotence ICV Admin. to ventromedial nucleus or medial preoptic area
(continued on next page)
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Table 5 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
MSH (␣)
Sexual behavior
Rat
Increased
Admin. to ventromedial nucleus or medial preoptic area
17
171
207
MSH (␣) MSH (␣) MSH/ACTH 4–10 analog Org2766 OT OT
Social interaction Social interaction Social interaction
Mouse Rat Rat
No change Decreased No change
2 2 13
1S 123 255 541
105 174 235
Human Rat
No change Preserved recognition
13 19
461 999
62 151
OT Sauvagine Sauvagine Sauvagine Somatostatin SP SP
Facial recognition test Social discrimination response Social recognition Aggressive behavior Defensive behavior Social interaction Aggressive behavior Aggressive behavior Aggressive behavior
Rat Mouse Mouse Mouse Fish Fish Mouse
Increased Decreased Increased Decreased Decreased Decreased Decreased
19 8 8 8 6 6 5
999 935 935 935 139 139 86
151 353 353 353 86 86 220
SP frag. (1–7) SP frag. (1–7)
Aggressive behavior Aggressive behavior
Mouse Mouse
Decreased Decreased
4 5
763 86
219 220
SP frag. (4–11)
Aggressive behavior
Mouse
No change
5
86
220
SP frag. analog pyroglutamylSP(7–11) SP frag. analog pyroglutamylSP(7–11) TRH
Aggressive behavior
Mouse
Increased
4
763
219
Aggressive behavior
Mouse
Increased
5
86
220
Dominance
Rat
Increased
9
539
441
TRH
Fighting
Rat
Decreased
9
539
441
TRH TRH analog RGH 2202 Tyr-MIF-1 Vasotocin VP
Sexual behavior RGH 2202 Sexual behavior Aggressive behavior Aggressive behavior Lordosis
Rat Rat
Increased Increased
12 12
1309 1309
152 152
Mouse Rat Hamster
Decreased Decreased Increased
7 5 14
1007 1135 1049
279 61 244
VP VP
Sexual behavior Social discrimination response Social interaction Social recognition Facial recognition test
Newt Rat
Increased Preserved recognition
4 19
97 999
365 151
Mouse Rat Human
No change Increased No change
2 19 13
1S 123 999 461
105 151 62
VP VP VP analog DGAVP
Admin. to central nucleus of amygdala Olfactory bulb admin. Olfactory bulb admin. ICV ICV ICV
Isolation-enhanced aggression Isolation-enhanced aggression; effect enhanced by naloxone Isolation-enhanced aggression
Isolation-enhanced aggression; effect antagonized by naloxone Admin. to nucleus accumbens septi; in dominant, but not subordinate, animals Admin. to nucleus accumbens septi; shockinduced and apomorphine-induced fighting
Pinealectomized animals Medial preoptic area admin. ICV or IP
Olfactory bulb admin.
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2213
Table 6 Learning and memory behaviors Peptide
Paradigm
Phase
Species
ACTH ACTH ACTH
Active avoidance Acquisition and recall Mouse Active avoidance Acquisition and recall Mouse Passive avoidance response Recall Rat
ACTH
Thomson Box visual discrimination Thomson Box visual discrimination Thomson Box visual discrimination Active avoidance
Effect
Notes
Vol Page
Ref
Acquisition
Rat
Improvement Given prenatally 9 Improvement Given postnatally 9 Decreased avoidance 10 latency No improvement 1
Extinction
Rat
No improvement
1
277
472
Reversal
Rat
No improvement
1
277
472
Extinction
Rat
Delayed
ICV
2
389
570
Passive avoidance response Acquisition
Rat
Improvement
ICV
2
389
570
Thomson Box visual discrimination Thomson Box visual discrimination Thomson Box visual discrimination Active avoidance
Acquisition
Rat
No improvement
1
277
472
Extinction
Rat
No improvement
1
277
472
Reversal
Rat
No improvement
1
277
472
Retention
Rat
Improvement
9
575
223
ACTH analog Ebiratide
Passive avoidance behavior Retention
Rat
Improvement
9
575
233
ACTH analog Ebiratide
Passive avoidance behavior Retention
Rat
Improvement
9
575
233
ACTH analog Ebiratide
Radial Maze (8 arm)
Working memory
Rat
Improvement
9
575
233
ACTH analog Ebiratide ACTH analog HOE 427
Retention Acquisition
Rat Rat
Improvement No change
9 10
575 303
233 456
Recall
Rat
No change
10
303
456
Reversal
Rat
Impaired reversal
10
303
456
ACTH analog ORG 2766 ACTH analog ORG 2766
Up-hill avoidance task Two-smell discrimination task Two-smell discrimination task Two-smell discrimination task Active avoidance Active avoidance
Acquisition and recall Mouse Acquisition and recall Mouse
Improvement Improvement
9 9
745 745
239 239
ACTH analog ORG 2766
Active behavior response
Extinction
Rat
Slowed
7
563
172
ACTH analog ORG 2766
Morris water maze
Recall
Rat
Improvement
11
911
432
ACTH analog ORG 2766
Passive avoidance response Recall
Rat
No change
11
911
432
ACTH analog ORG 2766 ACTH analog ORG 2766
Passive behavior response Passive behavior response
Acquisition Acquisition
Rat Rat
No change Improved
7 7
563 563
172 172
ACTH analog ORG 2766 ACTH analog ORG 2766
Passive behavior response Passive behavior response
Retention Retention
Rat Rat
Improved Improved
7 7
563 563
172 172
ACTH analog ORG 2766 ACTH analog ORG 2766 ACTH analog ORG 2766
Acquisition Extinction Acquisition
Rat Rat Rat
Improved No change No change
6 6 10
2S 41 2S 41 303
1 1 456
Recall
Rat
Improvement
10
303
456
ACTH/MSH frag. (4–10)
T-maze T-maze Two-smell discrimination task Two-smell discrimination task Active avoidance
Acquisition and recall Mouse
Improvement
745
239
ACTH/MSH frag. (4–10)
Active avoidance
Acquisition and recall Mouse
No change
745
239
ACTH ACTH ACTH analog -(Tyr9) melanotropin (9–18) ACTH analog -(Tyr9) melanotropin (9–18) ACTH analog D-PheMSH-ACTH ACTH analog D-PheMSH-ACTH ACTH analog D-PheMSH-ACTH ACTH analog Ebiratide
ACTH analog HOE 427 ACTH analog HOE 427
ACTH analog ORG 2766
Socially-deprived animals ECT-induced amnesia Scopolamineinduced amnesia Scopolamineinduced amnesia
Given prenatally Given postnatally; in males only Phase-shiftinduced amnesia Rats with fimbria fornix transection Rats with fimbria fornix transection Phase-shiftinduced amnesia Phase-shiftinduced amnesia 7–21 day old 7–21 day old
Given prenatally; 9 in females only Given postnatally 9
745 745 1117
239 239 425
277
472
(continued on next page)
2214
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 6 (continued) Peptide
Paradigm
ACTH/MSH frag. (4–10) ACTH/MSH frag. (4–10)
Benton visual retention test Cardiac orienting to novel stimuli Cardiac orienting to novel stimuli Classical conditioning Classical conditioning Conditioned taste aversion Digit symbol substitution test Discriminative learning
ACTH/MSH frag. (4–10) ACTH/MSH ACTH/MSH ACTH/MSH ACTH/MSH
frag. frag. frag. frag.
(4–10) (4–10) (4–10) (4–10)
ACTH/MSH frag. (4–10)
Phase
Species
Effect
Notes
Vol Page
Ref
Human Rabbit
Improvement Delayed
Elderly
Habituation
1 6
55 2S 97
359 228
Orienting
Rabbit
Enhanced
6
2S 97
228
Acquisition Extinction Retention
Rabbit Rabbit Rat Human
Worsening Faster Augmented version No change
3 3 1 1
715 715 207 55
478 478 501 359
Rat
Improvement
4
11
479
Rat
7
201
360
55 7
359 561
55 277
359 472
Discrimination stimulus Retention
Elderly In one out of ten rats
ACTH/MSH frag. (4–10)
Hunter delayed reaction apparatus
ACTH/MSH frag. (4–10) ACTH/MSH frag. (4–10)
Names and Faces test Passive avoidance response Retention
Human Rat
Enhanced at low doses inhibited at high doses No change Improvement
ACTH/MSH frag. (4–10) ACTH/MSH frag. (4–10)
Acquisition
Human Rat
No change Improvement
Extinction
Rat
Improvement
1
277
472
Reversal
Rat
No improvement
1
277
472
ACTH/MSH frag. (4–10) ACTH/MSH frag. (4–10) ACTH/MSH frag. (4–10)
Stroop test Thomson Box visual discrimination Thomson Box visual discrimination Thomson Box visual discrimination T-maze T-maze T-maze
Elderly 1 When admin. 3 prior to retention test, but not if admin. immediately after learning Elderly 1 1
Acquisition Extinction
Rat Rat Rat
7–21 day old 7–21 day old
6 6 10
2S 41 2S 41 1101
1 1 452
ACTH/MSH frag. (4–10) Actinomycin-D Amylin
Wechsler memory scale Passive avoidance Conditioned taste aversion
Elderly
Rat
1 13 13
55 927 961
359 308 79
Amylin Amylin
T-maze T-maze
Retention Retention
Human Rat No change Mouse Mouse
Improved No change Acted as a discrimination stimulus No change Impaired Intra-hypothalamic admin. Improvement Impairment
13 13
577 577
179 179
ANF, antiserum to ANF, antiserum to AT II
Active avoidance Extinction Passive avoidance response Consolidation Active avoidance
Rat Rat Rat
Facilitated Attenuated Improvement
13 13 21
957 957 407
46 46 37
AT II
Inhibitory shock avoidance
Retention
Rat
Impaired
16
1069
309
AT II AT II AT II
Passive avoidance Retention Passive avoidance behavior Passive avoidance behavior
Rat Rat Rat
Impaired Improvement No change
20 9 9
335 475 475
445 58 58
AT II AT II analog losartan
Plus maze Inhibitory shock avoidance
Rat Rat
Impaired Improved
20 16
335 1069
445 309
AT II analog saralasin BBS
Passive avoidance behavior Conditioned taste aversion Acquisition
Rat Rat
Improvement No change
Weak training ICV; strong training ICV ICV Hippocampal CA1 admin.; greater effect in right-side admin. Dentate gyrus admin. ICV ICV ICV with saralasin ICV Dentate gyrus admin.; AT IIinduced memory impairment ICV
9 2
475 61
58 301
ACTH/MSH frag. (4–10) ACTH/MSH frag. (4–10)
Retention
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2215
Table 6 (continued) Peptide
Paradigm
BBS
Species
Effect
Fixed interval responding
Rat
BBS BBS BBS Caerulein Caerulein
Non-reward operant task Reward operant task Taste aversion Active avoidance Morris water maze
Rat Rat Rat Rat Rat
Reduced responses after reinforcement Decreased Decreased No change Delayed Improvement
Caerulein
Morris water maze
Rat
Improvement
Caerulein
Morris water maze
Rat
Improved
Caerulein
Passive avoidance response Recall
Rat
Improved
Caerulein
Passive avoidance response Recall
Rat
Improved
Caerulein
Passive avoidance response Recall
Rat
Caerulein analog des-Gln-Caerulein Caerulein analog des-Gln-Caerulein Caerulein analog Leu-Nle-Caerulein Caerulein analog Leu-Nle-Caerulein Casein () analog D-TyrD-Phe3 CCK analog Boc-CCK-4 CCK analog Boc-CCK-4
Active avoidance response
Rat
Prevented ECSinduced amnesia Delayed
Extinction
Active avoidance response
Rat
Extinction
1435
55
4 4 3 10 13
1 1 61 843 1007
242 242 581 255 256
14
1073
535
12
699
537
11
1263
534
12
699
537
10
843
255
10
843
255
10
843
255
10
843
255
843
255
Amygdalakindled seizures
13
73
27
15 Central nucleus 19 of the amygdala admin.; post-trial injection 16 16
689 27
197 247
815 959
215 264
16 7 7 19 19 10 2 2 2 15
815 809 809 849 969 843 385 385 385 689
215 273 273 376 488 255 171 171 171 197
15
689
197
Rat Rat Rat
Increased Improved
Rat Rat
Rat Rat Rat Rat Human Rat Rat Rat Rat Rat
Impaired Potentiated amphetamine response Impaired No change Worsening No change Impaired recall Delayed Delayed Accelerated No change No effect
Rat
Increased
Extinction Acquisition Extinction Maintenance
12
10
Y-maze brightness discrimination Habituation Uphill avoidance task
Recall Acquisition Retention
Ref
ECS amnesia
Prevented ECSinduced amnesia Improved learning
Passive avoidance response Passive behavior response Passive behavior response Taste aversion conditioning Word recall Active avoidance response Conditioned feeding Conditioned feeding Conditioned feeding Conditioned place preference Habituation
Protein synthesis inhibitor-induced memory impairment Chronic ICV admin.; VIPinduced memory impairment Protein kinase-C inhibitor-induced memory impairment In rats with NMDA antagonistsinduced impairment. Protein kinase-C inhibitor-induced memory impairment ECS amnesia
Vol Page
ECS amnesia
Rat
Retention
Notes
Prevented ECSinduced amnesia Delayed
Passive avoidance response Recall
CCK CCK CCK CCK CCK CCK CCK CCK CCK CCK
CCK frag. CCK-8
Spatial memory
Rat
Active avoidance response Passive reward bar pressing
CCK-4 CCK-4 CCK-4 CCK-4 CCK-4 CCK-8 CCK-8 CCK-8 CCK-8 CCK-8
Extinction
Passive avoidance response Recall
CCK analog JMV-320 CCK analog L-365,260
frag. frag. frag. frag. frag. frag. frag. frag. frag. frag.
Phase
ICV ICV ICV, IV, or IP
(continued on next page)
2216
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 6 (continued) Peptide
Paradigm
CCK frag. CCK-8 CCK frag. CCK-8
Species
Effect
Non-reward operant task Passive avoidance response Recall
Rat Rat
CCK frag. CCK-8 CCK frag. CCK-8
Passive behavior response Passive behavior response
Acquisition Acquisition
Rat Rat
No change Prevented ECSinduced amnesia Improved Improved
CCK frag. CCK-8
Passive behavior response
Acquisition
Rat
Improved
CCK frag. CCK-8
Passive behavior response
Acquisition
Rat
Improved
CCK CCK CCK CCK CCK
Passive behavior response Retention Reward operant task Taste aversion conditioning Taste aversion conditioning Taste aversion conditioning Rat
Rat Rat Rat Rat Induced aversion Rat
Worsening Decreased Induced aversion ICV, IV, or IP Induced conditioning
frag. frag. frag. frag. frag.
CCK-8 CCK-8 CCK-8 CCK-8 CCK-8
Phase
CCK frag. CCK-8
Uphill avoidance task
CCK, antibody to
Discrimination task
CCK-8 CGRP, antiserum to CRF CRF
Free-operant avoidance Active avoidance response Active avoidance response Active avoidance response
CRF CRF
Arousal Conditioned taste aversion
CRF
CRF CRF binding protein
Fixed-interval food presentation Fixed-interval shock presentation Passive avoidance Shock-prod burying response Shock-prod burying test Active avoidance
CRF binding protein
Passive avoidance
Cycloheximide DDAVP Deltorphin
Passive avoidance Complex task execution Passive avoidance response Memory consolidation Fixed-interval food presentation Fixed-interval food presentation shock termination Delayed response in Working memory Wisconsin General Test apparatus Discrimination reversal Reversal Discrimination reversal Reversal task in Wisconsin General Test apparatus
CRF CRF CRF
Dermorphin Dermorphin
DSIP
DSIP DSIP
Retention
Notes
Vol Page
Ref
ECS amnesia
4 10
1 843
242 255
7 7
105 105
274 274
7
105
274
7
105
274
8 4 19 16 17
25 1 849 539 483
143 242 376 166 377
27
247
67
357
67 893 745 745
93 296 239 239
217 345
268 41
1199
384
ICV ICV; ECSinduced amnesia ICV; VPantiserum ECS-induced amnesia and VPantiserum
Central nucleus 19 of the amygdala admin.; post-trial injection Learning and recall Mouse Impairment Posterior 14 cingulate admin. Acquisition Rat Decreased avoidance 4 Extinction Rat Facilitated ICV 15 Acquisition and recall Mouse No change Given prenatally 9 Acquisition and recall Mouse Improvement in Given postnatally 9 females Monkey Increased ICV 4 Rat Induced aversion Third ventricle 20 admin. Monkey Increased responding ICV 10
Extinction
Improved
Monkey No change
ICV
10
1199
384
Rat Rat
Improved Decreased
ICV
13 13
927 1149
308 146
Rat Rat
Delayed Improvement
13 18
1149 711
146 227
Improved performance Rat Impaired Human Increased speed Mouse Increased step-down ICV latencies Monkey Increased responding ICV
18
711
227
13 12 11
927 1 591
308 76 421
10
1199
384
Monkey Decreased responding
10
1199
384
Monkey Worsening
2
1S 131 406
Monkey Worsening Monkey No change
2 2
1S 131 406 1S 131 406
Rat
ICV ICV; aged animals ICV
ICV
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2217
Table 6 (continued) Peptide
Paradigm
Dynorphin frag. (1–13)
Appetitive discrimination
Chicken Inhibited
Dynorphin frag. (1–13)
Avoidance conditioning
Chicken Inhibited
Endorphin ()
Act. behavior response
Recall
Rat
Endorphin ()
Acquisition
Endorphin ()
Fixed-ratio, fixed-interval food presentation Open field habituation
Monkey Decreased responding Rat Improved
Endorphin ()
Passive avoidance response Recall
Endorphin ()
Passive behavior response
Recall
Endorphin analog D-Ala2F5Phe4--endorphin
Delayed response in Wisconsin General Test apparatus Discrimination reversal task in Wisconsin General Test apparatus Discrimination reversal Conditioned aversion
Working memory
Reversed ECT8 induced amnesia but did not effect controls Rat Decreased avoidance 10 latency Rat Improved Reversed ECT8 induced- amnesia but did not affect controls Monkey Worsening 2
Reversal
Monkey No change
Reversal
Enkephalin analog D-Ala2- Delayed response in F5Phe4-Met-enkephalin Wisconsin General Test apparatus Enkephalin analog D-Ala2- Discrimination reversal F5Phe4-Met-enkephalin task in Wisconsin General Test apparatus Enkephalin analog D-Ala2-Met-enkephalinDelayed response in ethylamide Wisconsin General Test apparatus Enkephalin analog D-Ala2-Met-enkephalinDiscrimination reversal ethylamide task in Wisconsin General Test apparatus Enkephalin analog DPDPE Active avoidance response Enkephalin analog DPDPE Active avoidance response Enkephalin analog D-Phe4- Delayed response in Met-enkephalin Wisconsin General Test apparatus Enkephalin analog D-Phe4- Discrimination reversal Met-enkephalin task in Wisconsin General Test apparatus Enkephalin analog Fixed-ratio, fixed interval Metkephamide food presentation
Working memory
Monkey Worsening Rat Increased preference for conditioned stimulus, regardless of unconditioned stimulus Monkey Worsening
Reversal
Monkey No change
Working memory
Monkey Worsening
Reversal
Monkey No change
Acquisition Retention Working memory
Reversal
Endorphin analog D-Ala2F5Phe4--endorphin Enkephalin (D-Phe4-Met) Enkephalin (Met)
Phase
Species
Effect
Improved
Notes
Vol Page
Intermediate 14 medial hyperstriatum ventrale admin.; chicks Intermediate 14 medial hyperstriatum ventrale admin.; chicks Reversed ECT8 induced amnesia but did not affect controls 10
Ref
1165
95
1165
95
605
394
729
113
605
394
1117
425
605
394
1S 131 406
2
1S 131 406
2 In utero exposure 3
1S 131 406 881 520
2
1S 131 406
2
1S 131 406
2
1S 131 406
2
1S 131 406
Rat Impaired Rat Enhanced Monkey Worsening
13 13 2
885 340 885 340 1S 131 406
Monkey No change
2
1S 131 406
Rat
3
771
Decreased responding
305
(continued on next page)
2218
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 6 (continued) Peptide
Paradigm
Galanin Melanin
Species
Effect
Notes
Vol Page
Ref
Passive avoidance response Morris water maze Spatial memory
Mouse Rat
Decreased latency Improved
ICV In rats whose learning was inhibited by receiving MSG as neonates
16 12
1283 465
556 599
Melanin-concentrating hormone Melanin-concentrating hormone
Passive avoidance
Rat
Facilitated
15
757
343
Rat
Improved
20
1517
364
-(Tyr9) melanotropin(9–18) -(Tyr9) melanotropin(9–18) -(Tyr9) melanotropin(9–18) Melatonin
Active avoidance response
Extinction
Rat
Improved
Amygdala and hippocampus admin. ICV
7
11
540
T-maze
Reversal
Rat
Improved
ICV
7
11
540
T-maze
Spatial
Rat
Improved
ICV
7
11
540
Passive avoidance response Acquisition
Rat
Delayed
1
147
119
Melatonin
Passive avoidance response Extinction
Rat
Facilitated
Sham hypophysectomy but not hypophysectomy Sham hypophysectomy but not hypophysectomy
1
147
119
MIF-1
Delayed response in Wisconsin General Test apparatus Discrimination reversal task in Wisconsin General Test apparatus Lashly III maze Morris water maze
Working memory
Monkey Worsening
2
1S 131 406
Reversal
Monkey No change
2
1S 131 406
Learning Spatial memory
Mouse Rat
Improvement No change
11 12
527 465
Retention
Rat Rat
No change No change
2 8
1S 143 120 39 26
Retention
Rat
No change
8
39
26
Acquisition
Rat
Worsening
4
725
32
Reversal
Rat
No change
4
725
32
Extinction
Rat Rat Rat
Delayed No change No improvement
15 2 1
757 343 1S 143 120 277 472
Rat Improved Monkey Improved Rat Decreased errors
2 2 12
333 401 1S 131 406 929 450
MIF-1
MIF-1 MIF-1
MIF-1 MIF-1 analog Cyclo(Leu-Gly) MIF-1 analog Cyclo(Leu-Gly) MSH MSH MSH MSH MSH MSH MSH (␣) MSH (␣)
Phase
Extinction
Passive avoidance response
Passive avoidance response Conditioned avoidance response Conditioned avoidance response Fixed-ratio food presentation Fixed-ratio food presentation Passive avoidance Passive avoidance response Thomson Box visual discrimination Visual discrimination Discrimination reversal Visual discrimination learning
Acquisition
Reversal
Neonates In rats whose learning was inhibited by receiving MSG as neonates
Haloperidoltreated
Trend toward worsening
Medial anterior hypothalamic/ preoptic area admin.; no increase in responding to correct cues
115 599
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2219
Table 6 (continued) Peptide
Paradigm
Phase
MSH (␣)
Classically-conditioned fever response
MSH (␣)
Delayed response in Wisconsin General Test apparatus Discrimination reversal task in Wisconsin General Test apparatus Morris water maze
MSH (␣) MSH (␣)
MSH (␣)
Species
Effect
Notes
Vol Page
Ref
Classical conditioning Rat
11
1127
65
Working memory
Acted as unconditioned stimulus Monkey Improvement
2
1S 131 406
Reversal
Monkey No change
2
1S 131 406
Spatial memory
Rat
Improved
12
465
599
Acquisition
Rat
No change
4
145
222
Reversal
Rat
No change
4
145
222
Acquisition
Rat
Improved
4
145
222
Reversal
Rat
Improved
4
145
222
Acquisition
Rat
Facilitated
Hypophysectomy, 1 but not in controls Hypophysectomy, 1 but not control ECS amnesia 10
147
119
147
119
361
36
10 1
361 277
36 472
1
277
472
4 4 4 4 4 4 4 2
721 721 721 145 145 145 145 101
286 286 286 222 222 222 222 402
In rats whose learning was inhibited by receiving MSG as neonates
MSH (␣)
Operant auditory discrimination Operant auditory discrimination Operant visual discrimination Operant visual discrimination Passive avoidance response
MSH (␣)
Passive avoidance response Extinction
Rat
Delayed
MSH (␣)
Passive avoidance response Recall
Rat
MSH (␣) MSH (␣)
Passive avoidance response Thomson Box visual discrimination Thomson Box visual discrimination Y maze Y maze Y maze Y maze Y maze Y maze Y maze Y maze visual discrimination test Y maze visual discrimination test Y maze visual discrimination test Y maze
Recall Extinction
Rat Rat
Prevented ECSinduced amnesia No change No change
Reversal
Rat
Improvement
Acquisition Extinction Reversal Acquisition Acquisition Reversal Reversal Acquisition
Rat Rat Rat Rat Rat Rat Rat Rat
Improvement No change Improvement Improved No change Improved No change Improvement
Extinction
Rat
No change
2
101
402
Reversal
Rat
Improvement
2
101
402
Acquisition
Rat
No change
ICV
4
721
286
Y maze
Extinction
Rat
No change
ICV
4
721
286
Y maze
Reversal
Rat
Worsening
ICV
4
721
286
Passive avoidance response Recall
Rat
No change
ECS amnesia
10
361
36
Passive avoidance response Recall
Rat
Improved recall
With controls but 10 not ECS amnesia
361
36
MSH (␣) MSH (␣) MSH (␣)
MSH (␣) MSH MSH MSH MSH MSH MSH MSH MSH
(␣) (␣) (␣) (␣) (␣) (␣) (␣) (␣)
MSH (␣) MSH (␣) MSH analog [Nle,D-Phe]␣-MSH MSH analog [Nle,D-Phe]␣-MSH MSH analog [Nle,D-Phe]␣-MSH MSH analog Ac-Cys-GluHis-D-Phe-Arg-Trp-CysNH2 MSH analog Ac-Cys-GluHis-Phe-Arg-Trp-CysNH2
ICV ICV ICV
(continued on next page)
2220
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 6 (continued) Peptide
Paradigm
MSH analog Ac-Nle-AspHis-D-Phe-Arg-Trp-LysNH2 MSH analog Ac-Nle-GluHis-D-Phe-Arg-TrpDab-NH2 MSH analog Ac-Nle-GluHis-D-Phe-Arg-Trp-GlyLys-NH2 MSH analog Ac-Nle-GluHis-D-Phe-Arg-Trp-GlyLys-NH2 MSH analog Ac-Nle-GluHis-D-Phe-Arg-Trp-GlyLys-NH2 MSH analog Ac-Nle-GluHis-D-Phe-Arg-Trp-GlyLys-NH2 MSH analog Ac-Nle-GluHis-D-Phe-Arg-Trp-GlyNH2 MSH analog Ac-Nle-GluHis-D-Phe-Arg-Trp-GlyNH2 MSH analog Ac-Nle-GluHis-D-Phe-Arg-Trp-GlyOH MSH analog H-Cys-GluD-His-Phe-D-Arg-TrpD-Cys-OH MSH analog H-Cys-GluHis-Phe-D-Arg-Phe-DCys-OH MSH analog H-Cys-GluHis-Phe-D-Arg-Phe-DCys-OH MSH analog H-Cys-GluHis-Phe-D-Arg-Phe-DCys-OH MSH analog H-Cys-GluHis-Phe-D-Arg-Trp-CysOH MSH analog H-Cys-GluHis-Phe-D-Lys-Phe-CysOH MSH analog H-Cys-GluHis-Phe-D-Lys-Phe-DCys-OH MSH analog MK-771 MSH () MSH () MSH () MSH (␥) MSH (␥)
Phase
Species
Effect
Notes
Vol Page
Ref
Passive avoidance response Recall
Rat
No change
ECS amnesia
10
361
36
Passive avoidance response Recall
Rat
Prevented ECSinduced amnesia
ECS amnesia
10
361
36
Passive avoidance response Recall
Rat
Prevented ECSinduced amnesia
ECS amnesia
10
361
36
Thomson-Bryant discrimination test
Acquisition
Rat
No change
10
361
36
Thomson-Bryant discrimination test
Retention
Rat
No change
10
361
36
Thomson-Bryant discrimination test
Reversal
Rat
Impaired reversal
10
361
36
Passive avoidance response Recall
Rat
Prevented ECSinduced amnesia
10
361
36
Passive avoidance response Recall
Rat
No change
10
361
36
Passive avoidance response Recall
Rat
Prevented ECSinduced amnesia
ECS amnesia
10
361
36
Passive avoidance response Recall
Rat
Prevented ECSinduced amnesia
ECS amnesia
10
361
36
Thomson-Bryant discrimination test
Acquisition
Rat
No change
10
361
36
Thomson-Bryant discrimination test
Retention
Rat
No change
10
361
36
Thomson-Bryant discrimination test
Reversal
Rat
No change
10
361
36
Passive avoidance response Recall
Rat
No change
ECS amnesia
10
361
36
Passive avoidance response Recall
Rat
No change
ECS amnesia
10
361
36
Passive avoidance response Recall
Rat
No change
ECS amnesia
10
361
36
Fixed-ratio, fixed-interval shock avoidance Thomson Box visual discrimination Thomson Box visual discrimination Thomson Box visual discrimination Y maze visual discrimination test Y maze visual discrimination test
Monkey Increased
ECS amnesia
Acquisition
Rat
No Improvement
Under fixed-ratio 5 schedule 1
783
56
277
472
Extinction
Rat
No improvement
1
277
472
Reversal
Rat
Improvement
1
277
472
Acquisition
Rat
No change
2
101
402
Extinction
Rat
No change
2
101
402
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2221
Table 6 (continued) Peptide
Paradigm
Phase
Species
Effect
MSH (␥)
Reversal
Rat Rat
NPY
Y maze visual discrimination test Progressive-ratio operant schedule T-maze
Retention
Mouse
NPY
T-maze
Retention
Mouse
NPY frag. (20–36)
T-maze
Retention
Mouse
NPY frag. (20–36)
T-maze
Retention
Mouse
NPY frag. (26–36)
T-maze
Retention
Mouse
NT
Rewarded behavior
Rat
OT
Buschke restrictive reminding test Escape behavior
Human
Facial recognition test Memory comparison test One-trail step-through Retention paradigm Passive avoidance response Learning/recall Social discrimination response Social recognition
Human Human Rat
Stroop color-word test Y-maze brightness discrimination Passive avoidance T-maze T-maze
Recall
Human Rat
Retention Retention
Rat Mouse Mouse
Pancreastatin frag. (33–49) T-maze Retention Secretin Novel-object approach Secretin Passive avoidance Somatostatin Passive avoidance response
Mouse Rat Rat Rat
Somatostatin
Passive avoidance response Recall
Rat
Somatostatin (7–10) Somatostatin frag. (3–8) Somatostatin frag. (9–14) SP
Passive avoidance response Recall Passive avoidance response Recall Passive avoidance response Recall Morris water maze
Rat Rat Rat Rat
SP
Tunnel maze
Short and long term memory
Rat
SP SP analog [pGlu6]SP(6–11) SP frag. (1–7) TRH
Up-hill avoidance learning Up-hill avoidance learning
Retention Retention
Up-hill avoidance learning Active avoidance response
Retention
NPY
OT
OT OT OT OT OT OT OT OT PACAP Pancreastatin Pancreastatin
Vol Page
Ref
Worsening
2
101
402
Increased responding Perifornical region admin. Improvement ICV; weak training Impairment ICV; strong training Improvement ICV; weak training Impairment ICV; strong training No change ICV; weak training Decreased ICV responding Reduced initial storage Increased With central, but not peripheral, admin. No change No change No change
19
1667
60
10
963
177
10
963
177
10
963
177
10
963
177
10
963
177
16
1417
457
13
461
62
2
1
350
13 13 15
461 461 229
62 62 52
927 999
529 151
999
151
461 717
62 84
1067 1077 1077
536 180 180
1077 739 1067 293
180 83 536 568
1153
569
1153 1153 1153 275
569 569 569 504
85
385
Rat Rat
Prenatal exposure 11 Olfactory bulb 19 admin. Olfactory bulb 19 admin. No change 13 No change Newborns and 9 adults No change 14 Improvement 9 Improvement Scopolamine9 induced amnesia Improvement 9 Increased ICV 4 No change 14 Blocked ECTICV; ECT4 induced amnesia induced amnesia Decreased avoidance ICV 10 latency No change ICV 10 No change ICV 10 No change ICV 10 Improved Rats with lesions 17 performance of the hippocampus with or without fetal grafts Facilitated With pretrial, but 14 not post-trial, admin. Improvement 11 No change 11
163 163
224 224
Rat Rat
Improvement Improved
163 1309
224 152
Rat
Rat Rat Rat
Notes
No change Preserved recognition Increased
Rats with hypoxic amnesia
11 12
(continued on next page)
2222
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 6 (continued) Peptide
Paradigm
TRH
Active avoidance response
TRH
TRH analog MK-771
Fixed-ratio, fixed-interval food presentation Fixed-ratio, fixed-interval food presentation Fixed-ratio, fixed-interval food presentation Fixed-ratio, fixed-interval shock avoidance Passive avoidance response Radial arm maze Fixed-ratio, fixed-interval food presentation Fixed-ratio, fixed-interval food presentation Fixed-ratio, fixed-interval food presentation Radial maze (12-arm)
TRH analog RGH 2202
Active avoidance response
Recall
TRH analog RGH 2202
Active avoidance response
Recall
TRH analog RGH 2202 TRH analog RGH 2202 TSH
Passive avoidance response Recall Radial arm maze Spatial memory Fixed-interval food reward with fixed-ratio shock Fixed-interval food reward with fixed-ratio shock
TRH TRH TRH TRH TRH TRH analog MK-771 TRH analog MK-771 TRH analog MK-771
TSH
Tuftsin
Phase
Species
Effect
Rat
Monkey Recall Spatial memory
Working memory
Reversed age-related Aged animals impairment Monkey Decreased responding Pigeon Decreased responding Rabbit Decreased responding Increased Under fixed-ratio schedule Rat Improvement Rat Decreased errors Monkey Decreased responding Pigeon Decreased responding Rabbit Decreased responding Rat Improvement In medal septal lesioned but not controls Rat Reversed hypoxiaRats with induced amnesia hypoxic amnesia Rat Reversed age-related Aged animals impairment Rat Improvement Rat Decreased errors Pigeon Decreased When admin. alone Pigeon Increased When admin. with pentobarbital, chlordiazepoxide, or alcohol Monkey Increased responding ICV
Urocortin
Fixed-interval food presentation Fixed-interval shock presentation Conditioned taste aversion
Vasotocin
Passive avoidance behavior Retention
Chicken Increased
Vasotocin
Acquisition
Rat
No change
Reversal
Rat
Improvement
VIP
Thompson-Bryant discrimination box Thompson-Bryant discrimination box Morris water maze
Rat
Impairment
VIP VIP VIP
Passive avoidance Passive avoidance T-maze
Rat Rat Mouse
Impairment No change Impairment
VP VP
Active behavior response Cognition
Rat Human
Slowed Improved
Tuftsin
Vasotocin
Notes
Monkey Decreased responding Rat Induced aversion
Retention
Extinction Activation, alertness, and fatigue
ICV
Vol Page
Ref
12
1309
152
4
177
23
4
177
23
4
177
23
5
783
56
12 12 4
1309 1309 177
152 152 23
4
177
23
4
177
23
10
121
240
12
1309
152
12
1309
152
12 12 5
1309 1309 809
152 152 591
5
809
591
10
1199
384
10
1199
384
345
41
1221
122
383
35
6
383
35
14
1073
535
14 14 11
1067 1067 933
536 536 176
7 12
213 1385
290 429
Third ventricle 20 admin. ICV or IP; 2 day 5 old chicks 6
Chronic ICV admin. Central admin. Peripheral admin. ICV; effect antagonized by arecoline, naloxone and ST 587, but not CCK or NPY ICV
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2223
Table 6 (continued) Peptide
Paradigm
Phase
Species
Effect
Notes
Vol Page
Ref
VP
Conditioned aversion response
Acquisition
Rat
ICV
7
213
290
VP
Escape behavior
Rat
Did not act as unconditioned stimulus Increased
2
1
350
VP
Habituation
Mouse
Increased
With central, but not peripheral, admin. Transgenic admin.
16
1329
362
VP VP
Recall Recall
Human Human
Enhanced free recall No change
12 12
1379 1379
391 391
Retention
Rat
Impairment
15
229
52
Human
No change
12
1393
20
VP
Incidental memory task Oddball paradigm word recall One-trail step-through paradigm Orienting response and stimulus mismatch Passive avoidance
Mouse
No change
1329
362
VP VP
Passive avoidance behavior Recall Passive avoidance response Learning/recall
Rat Rat
493 927
550 529
VP VP VP
Passive avoidance response Retention Passive behavior response Acquisition Passive behavior response Acquisition
Rat Rat Rat
Impairment Increased step-down latency (improvement) in females Improvement No change Improved
Transgenic 16 admin. Given prenatally 8 Prenatal exposure 11
11 7 7
633 105 105
317 274 274
VP VP
Radial maze (12 arm) Social discrimination response Social recognition
Working
Rat Rat
1 19
261 999
66 151
19
999
151
Thompson-Bryant discrimination box Thompson-Bryant discrimination box Thompson-Bryant discrimination test Thompson-Bryant discrimination test Thompson-Bryant discrimination test Thomson-Bryant discrimination test Thomson-Bryant discrimination test Conditioned avoidance for smoking cessation Conditioned avoidance for smoking cessation Emotionally-influenced free recall Impaired free recall performance Radial maze (24-arm)
VP VP
VP VP VP VP VP VP VP VP VP (Lys) VP (Lys) VP (Lys) VP (Lys) VP analog 8-D-arginineVP VP analog 8-L-argininedeamino-VP VP analog DAVP VP analog DDAVP
Neonatal exposure
ICV ICV; ECSinduced amnesia
Rat
No change Preserved recognition Increased
Acquisition
Rat
No change
6
383
35
Reversal
Rat
Improvement
6
383
35
Acquisition
Rat
No change
Given prenatally
8
493
550
Reversal
Rat
No change
Given prenatally
8
493
550
Reversal recall
Rat
Impairment
Given prenatally
8
493
550
Acquisition
Mouse
No change
10
237
549
Reversal
Mouse
Improvement
237
549
Acquisition
Human
Improvement
In albino, but not 10 hooded, rats 3
527
164
Extinction
Human
Accelerated
3
527
164
Human
No change
Twins
6
297
170
Human
No change
Twins
6
297
170
Working memory
Rat
No change
3
725
67
Radial maze (24-arm)
Working memory
Rat
No change
3
725
67
Radial maze (12-arm) Active avoidance
Working Acquisition
Rat Rat
No change Enhanced at low dose and decreased at high dose
1 12
261 471
66 493
Olfactory bulb admin. Olfactory bulb admin.
(continued on next page)
2224
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 6 (continued) Peptide
Paradigm
Phase
Species
Effect
Notes
Vol Page
Ref
VP analog DDAVP
Active avoidance
Extinction
Rat
Inhibition
Dose-dependent effect
12
471
493
VP VP VP VP
Active avoidance test Active avoidance test Benton visual retention test Color and shape discrimination Delayed word list recall Immediate recall of sentences Immediate word list recall Implicational sentence recall
Acquisition Extinction
Rat Rat Human Human
Improved Delayed No change Improvement
6 6 3 3
23 23 627 627
221 221 29 29
Recall
Human Human
No change Improved
11 6
1313 397
31 548
Recall Recall
Human Human
11 11
1313 473
31 33
11 11 7 7
1313 633 563 563
31 317 172 172
7 7
563 563
172 172
16 1 5
1327 261 819
30 66 34
4
707
28
analog analog analog analog
DDAVP DDAVP DDAVP DDAVP
VP analog DDAVP VP analog DDAVP VP analog DDAVP VP analog DDAVP
Males Males
Paper folding test Passive avoidance response Retention Passive behavior response Acquisition Passive behavior response Acquisition
Human Rat Rat Rat
Impaired Dose-dependent impairment or facilitation No change Improvement No change Improved
VP analog DDAVP VP analog DDAVP
Passive behavior response Passive behavior response
Acquisition Acquisition
Rat Rat
Improved Improved
VP analog DDAVP VP analog DDAVP VP analog DDAVP
Phrase recall Radial maze (12-arm) Sentence comprehension and recall
Working
Human Rat Human
No change Worsening Improvement
VP analog DDAVP
Human
Improved attention
Acquisition
Human Rat
No change Improvement
11 3
1313 521
31 100
Extinction
Rat
No change
3
521
100
Reversal
Rat
Impairment
3
521
100
VP analog DDAVP
Sternberg item recognition task Stroop color word test Thomson Box visual discrimination Thomson Box visual discrimination Thomson Box visual discrimination Word list recall
Human
9
1361
430
VP analog DDAVP
Word recall
Human
Improved recent recall, worsened distant recall Improved
16
179
63
VP analog DDAVP
Y-maze brightness discrimination Y-maze brightness discrimination Classical conditioning
Acquisition
Rat
Improvement
9
717
84
Recall
Rat
Improvement
9
717
84
Acquisition
Rabbit
No change
4
37
229
VP analog Deaminodicarba-arginine-8-VP
Classical conditioning
Acquisition
Rabbit
No change
4
37
229
VP analog Deaminodicarba-arginine-8-VP
Classical conditioning
Extinction
Rabbit
Accelerated
4
37
229
VP analog Deaminodicarba-arginine-8-VP
Classical conditioning
Extinction
Rabbit
No change
4
37
229
VP VP VP VP
analog analog analog analog
DDAVP DDAVP DDAVP DDAVP
VP analog DDAVP VP analog DDAVP VP analog DDAVP VP analog DDAVP
VP analog DDAVP VP analog Deaminodicarba-arginine-8-VP
Phase-shiftinduced amnesia Phase-shiftinduced amnesia
Effect seen in males but not females Males
In men but not women Newborns but not adults Newborns and adults Classicallyconditioned bradycardia Classicallyconditioned nictitating membrane response Classicallyconditioned bradycardia Classicallyconditioned nictitating membrane response
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2225
Table 6 (continued) Peptide
Paradigm
Phase
Species
Effect
VP analog desGlyNH2DDAVP VP analog desGly-NH2VP VP analog DGAVP VP analog DGAVP
Radial maze (24-arm)
Working memory
Rat
Radial maze (12-arm)
Working
Active behavior response Active behavior response
Extinction Extinction
VP analog DGAVP
Buschke restrictive reminding test
VP analog DGAVP
Cardiac orienting to novel stimuli Cardiac orienting to novel stimuli Facial recognition test Memory comparison test
VP analog DGAVP VP analog DGAVP VP analog DGAVP
Notes
Vol Page
Ref
Improvement
3
725
67
Rat
No change
1
261
66
Rat Rat
Slowed Slowed
7 7
563 563
172 172
Human
13
461
62
6
2S 97
228
Phase-shiftinduced amnesia
Habituation
Rabbit
Increased word recall and delayed recognition Delayed
Orienting
Rabbit
Enhanced
6
2S 97
228
Human Human
13 13
461 461
62 62
13
461
62
7 11 11
213 633 633
290 317 317
Stroop color-word test
Human
No change Diminished reaction time No change
Active behavior response Extinction Passive avoidance response Retention Passive avoidance response Retention
Rat Rat Rat
Faster Improvement Improvement
Thompson-Bryant Acquisition discrimination box Thompson-Bryant Reversal discrimination box Passive avoidance response Retention
Rat
No change
6
383
35
Rat
No change
6
383
35
Rat
Improvement
11
633
317
Passive avoidance response Retention
Rat
Improvement
11
633
317
Passive avoidance response Retention Passive avoidance behavior Retention
Rat No change Chicken Enhanced avoidance
11 4
633 401
317 123
Passive avoidance response Post-learning application
Rat
No change
6
2S 177 566
VP, antibody to
Passive avoidance response Retention
Rat
Impairment
6
2S 177 566
VP, antibody to VP, antibody to
Passive avoidance response Retention Passive avoidance response Retention
Rat Mouse
Impairment Retention deficit
6 4
2S 177 566 265 306
VP analog DGAVP VP analog dPTyr(me) ArgVP VP analog D-ZNC(C)PR VP analog D-ZNC (C-OMe)PR VP analog Pressinoic acid VP analog Pressinoic acid VP analog ZNC(C)PR (VP 4–8) VP analog ZNC(COMe)PR VP analog ZNCPR dimer VP frag. L-prolyl-Larginyl-glycinamide VP, antibody to
ICV; VP treated
ICV but not IP; 2 day old chicks Dorsal septum admin. after learning Hippocampal admin. ICV ICV
2226
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 7 Stereotyped and other behaviors sorted by peptide Peptide
Behavior
Species
Effect
ACTH ACTH ACTH ACTH
Distress vocalization Grooming Grooming Grooming
Chicken Rat Rat Rat
Increased Increased Increased Increased
ACTH ACTH
Grooming Grooming
Rat Rat
Increased Increased
ACTH
Grooming
Rat
Increased
ACTH
Grooming
Rat and mouse
Increased
ACTH
Mouse
No change
ACTH ACTH
Myoclonus (serotonergic behavior) Vomiting Yawning
Dog Rat
No change Increased
ACTH frag. (1–24) ACTH frag. (1–24)
Anxiety behaviors Grooming
Chicken Rat
Increased Increased
ACTH frag. (1–4)
Grooming
Rat
Enhanced
ACTH frag. (4–10) ACTH frag. (4–10) ACTH/MSH 4–10 analog ORG 2766 ACTH/MSH frag. analogs, C-frag. elongated or cyclized Adrenal Peptide E (Frog)
Anxiety behaviors Distress vocalization Rotation
Chicken Chicken Rat
Increased Increased Decreased
Grooming
Rat
Increased
Mouse
No change
Rat Dog Rat
BBS BBS BBS
Morphine-induced behaviors Grooming Vomiting Amphetamine-induced stereotypy Apomorphine-induced stereotypy Bite/scratch behavior Grooming Grooming
Vol
Page
Ref
11 7 4 5
915 597 907 713
414 476 202 546
6 7
369 651
562 160
16
1263
434
8
841
159
10
5
439
6 16
1S 173 1263
597 434
11 4
915 833
414 6
7
1
134
11 11 14
915 915 1317
414 414 10
5
1197
232
ICV
17
1291
97
No change No change Increased
ICV ICV ICV
17 6 9
589 1S 173 475
91 597 58
Rat
Increased
ICV
9
475
58
Mouse Rat Rat
Increased Increased Increased
7 2 8
835 2S 179 237
47 198 303
Grooming Grooming Grooming
Rat Rat Rat
Increased Increased Increased
12 13 4
761 1215 693
182 99 355
BBS
Grooming
Rat
Increased
7
557
352
BBS
Grooming
Rat
Increased
9
1S 245
260
BBS
Grooming
Rat
Increased
9
1S 245
260
BBS
Grooming
Rat
Increased
Intraspinal admin. ICV With admin. to multiple brain areas ICV ICV; developed over 8 days ICV; effect antagonized by neuroleptics ICV; antagonized by morphine With admin. to nucleus tractus solitarius but not nucleus accumbens With admin. to nucleus tractus solitarius but not nucleus accumbens With ICV but not IV admin.; no antagonism by haloperidol, morphine, naloxone, or NT. No effect of hypophysectomy or adrenalectomy
BBS BBS BBS
4
907
202
Amylin AT AT II AT II
Notes ICV; blocked by naloxone ICV Pre-optic, anterior hypothalamic admin. ICV ICV; area of action localized to anterior-ventral third ventricle ICV; not antagonized by nitric oxide synthase inhibitor ICV; greater effect in rats than mice 5-hydroxytryptophaninduced ICV ICV; antagonized by nitric oxide synthase inhibitor ICV; effect antagonized by naloxone In MSH-induced grooming; no effect alone
Nigrostriatal lesions and amphetamine-induced turning More potent than unmodified frag.
(continued on next page)
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2227
Table 7 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
BBS
Grooming
Rat
Increased
6
369
562
BBS
Grooming
Rat
Increased
6
1179
564
BBS
Grooming
Rat
Increased
6
1179
563
BBS BBS BBS BBS BBS
Grooming and scratching Licking Licking Respiration Righting
Rat Rat Rat Rat Rat
Increased Decreased Decreased Increased Impaired
ICV; different doses induce different types of grooming Effect enhanced by haloperidol, antagonized by naloxone and NT ICV; antagonized by haloperidol Neonates
10 17 16 4 2
529 107 903 277 1S 99
259 175 523 395 187
BBS
Scratching
Mouse
Increased
9
909
567
BBS
Scratching
Rat
Increased
6
1179
563
BBS BBS Bradykinin Calcitonin Calcitonin
Sighing Vomiting Vomiting Respiration Writhing
Rat Dog Dog Rat Mouse
Increased No change No change Increased Increased analgesia
4 6 6 4 6
277 1S 173 1S 173 277 3S 277
395 597 597 395 200
Casomorphin analog BCH 325 Casomorphin analog BCH 325
Jerking
Rat
Decreased
16
635
447
Yawning
Rat
Normalized
16
635
447
Casomorphin analogs
Yawning
Rat
Decreased
13
69
464
Casomorphin frag. (1–5) CCK CCK CCK
Distress vocalization Anxiety behavior Respiration Stretching
Chicken Rat Cat Mouse
Decreased Increased Increased tidal volume Decreased
5 16 9 10
829 1313 809 31
415 263 246 9
CCK analog Boc-CCK-4
Anxiety behavior
Rat
No change
19
27
247
CCK frag. CCK-8
Anxiety behavior
Rat
No change
19
27
247
CCK-4 CCK-4 CCK-8
Circling Defecation Catalepsy
Rat Rat Rat
No change No change Blocked
7 6 6
809 91 237
367 275 253
CCK-8 CCK-8 CCK-8
Circling Defecation Rotation
Rat Rat Rat
Increased No change Increased
7 6 18
809 91 1161
367 275 396
CCK-8 CCK-8, desulfated Ceruletide
Rumination Circling Catalepsy
Sheep Rat Rat
No change No change Decreased
5 7 10
81 809 779
238 367 250
Ceruletide analogs Ceruletide analogs
Catalepsy Convulsions
Mouse Mouse
Increased Decreased
3 3
701 701
610 610
ICV ICV; alcohol-induced righting impairment Intrathecal admin.; not antagonized by D-Pro2,DTrp7,9-SP and phenoxybenzamine ICV; antagonized by naloxone ICV ICV ICV ICV ICV, IP, and IV, but no effect SC; Ach-induced writhing Bromocriptine-induced jerks Bromocriptine-evoked yawning that was inhibited by reserpine ICV; apomorphine-induced yawning ICV; chicks Ventral medulla admin. ICV; abdominal irritantinduced stretching Central nucleus of the amygdala admin. Central nucleus of the amygdala admin. ICV Nucleus accumbens admin. ICV; endorphin-induced catalepsy; effect blocked by frontal cortex lesion ICV Nucleus accumbens admin. Animals pre-treated with peptide prior to nigrostriatal lesions; dopaminedepletion-induced turning ICV Haloperidol-induced catalepsy Hexobarbital-induced sleep Harman-induced convulsions
(continued on next page)
2228
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 7 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
Ceruletide analogs
Gnawing
Mouse
Increased
Methylphen-induced gnawing
2
2S 65
609
Ceruletide analogs CGRP
Ptosis Grooming
Mouse Rat
Increased Increased
2 17
2S 65 1183
609 294
CGRP CRF
Grooming Boxing behavior
Rat Rat
Decreased Decreased
5 6
861 891
297 107
CRF CRF CRF
Grooming Grooming Grooming
Rat Rat Rat
Increased Increased Increased
13 6 20
1149 3S 53 509
146 392 571
CRF
Grooming
Rat and mouse
Increased
8
841
159
CRF CRF antagonist ␣-helical CRF Cyclo (Leu-Gly)
Huddling and lying down Freezing behavior
Monkey Rat
Increased Decreased
4 15
217 1303
268 237
Rat
Decreased
8
39
26
Cyclo (Leu-Gly)
Apomorphine-induced stereotypy Rotation
Rat
No change
8
39
26
Dermorphin
Catalepsy
Rat
Increased
6
3S 165
59
Dermorphin
Catalepsy
Rat
Decreased
6
3S 165
59
Dynorphin analog desTyr1-dynorphin (1–13) Dynorphin frag (1–13) Dynorphin frag. (1–13)
Grooming
Rat
Increased
4
833
6
Barrel rolling Grooming
Rat Rat
Increased Increased
8 4
1089 833
201 6
Dynorphin frag. (1–13)
Grooming
Rat and mouse
Increased
1
341
578
Dynorphin Dynorphin Dynorphin Dynorphin Dynorphin
Hind limb paralysis Shaking Circling Circling Circling
Rat Rat Rat Rat Rat
Increased Increased Increased Increased Increased
4 8 8 8 8
631 1089 837 837 837
168 201 186 186 186
Dynorphin frag. (2–17) Endorphin (␣)
Circling Distress vocalization
Rat Chicken
Increased Decreased
8 5
837 823
186 573
Endorphin ()
Catalepsy
Rat
Increased
6
237
253
Endorphin ()
Distress vocalization
Chicken
Decreased
5
823
573
Endorphin ()
Righting
Rat
Impaired
2
1S 99
187
Endorphin () Endorphin ()
Startle Trembling
Rat Rat
Decreased Decreased
2 2
1S 137 1S 99
234 187
Endorphin (␥)
Amphetamine-induced stereotypy
Rat
Decreased
2
9
285
Endorphin (␣)
Distress vocalization
Chicken
Decreased
5
823
573
Enkephalin
Vocalization
Sheep
Decreased
9
1221
417
frag. frag. frag. frag. frag.
(1–13) (1–13) (1–6) (1–7) (1–8)
Antagonized by nitric oxide synthase inhibitor ICV Medial frontal cortex admin.; carbachol-induced boxing behavior ICV ICV but not SC ICV; not antagonized by melanocortin-4 receptor antagonist ICV; greater effect in rats than mice; not antagonized by dexamethasone ICV Foot-shock-induced behavior Rats treated chronically with haloperidol Apomorphine-induced rotation in rats with unilateral nigrostriatal lesions ICV; effects increased by caudate lesion ICV; in dermorphin-tolerant but not morphine-tolerant rats ICV; effect antagonized by naloxone ICV or IV ICV; effect antagonized by naloxone ICV; not reversible by naloxone Intrathecal admin. ICV or IV Substantia nigra admin. Substantia nigra admin. Substantia nigra admin.; not antagonized by naloxone Substantia nigra admin. With ICV, but not IP, admin. ICV; effects enhanced by frontal cortex ablation With ICV, but not IP, admin.; chicks ICV; alcohol-induced righting impairment ICV; alcohol-induced tremors Effect antagonized by 2bromo-ergocryptine and naloxone With ICV, but not IP, admin. ICV
(continued on next page)
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2229
Table 7 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
Enkephalin (Leu) Enkephalin (Met) Enkephalin analog DADLE
Circling Shaking Respiration
Rat Fish Rat
Increased Increased Increased
Substantia nigra admin.
8 6 3
837 139 1031
186 86 225
Enkephalin analog DADLE Enkephalin analog DAGO
Respiration Respiration
Rat Rat
3 3
1023 1031
236 225
Enkephalin analog DAMA Enkephalin analog FK33– 824 Galanin Galanin
Shaking Respiration
Rat Mouse
Increased Increased at low dose decreased at high dose Increased Decreased rate
4 9
239 777
155 378
Anxiety behavior Freezing behavior
Mouse Rat
No change No change
16 15
1283 1303
556 237
Galanin
Grooming
Rat
No change
11
1001
304
Galanin
Rat
No change
15
1303
237
Gastrin (1–17) Gastrin-releasing peptide Gastrin-releasing peptide
Opiate withdrawal behavior Rumination Bite/scratch behavior Licking
Sheep Mouse Rat
Increased Increased Decreased
5 7 17
81 835 107
238 47 175
Gastrin-releasing peptide Kassinin L-pyroglutamyl-L-leucine
Licking Bite/scratch behavior Jumping
Rat Mouse Rat
Decreased Increased Delayed
16 7 4
903 835 417
523 47 295
MCH
Anxiety behavior
Rat
Decreased
17
171
207
Melatonin MIF-1
Startle response Catalepsy
Rat Mouse
Decreased Prevented tolerance
2 8
1S 155 1051
118 380
MIF-1
Catalepsy
Rat
Decreased
2
105
85
MIF-1
Circling
Rat
No change
2
189
148
MIF-1 MIF-1
Defecation Head turning
Rat Rat
Decreased No change
2 2
1S 143 189
120 148
MIF-1 MIF-1
Head twitch Oral act. (serotonergic behavior)
Mouse Rat
No change No change
2 14
189 1159
148 206
MIF-1 MIF-1 analog 3(R)-[(2(S)pyrrolidinylcarbonyl) amino]-2-oxo-1pyrrolidineacetamide MIF-1 analog Cyclo(Leu-Gly) MIF-1 analog parepide
Depressive behaviors Rotation
Human Rat
Decreased Increased
4 18
297 1209
559 363
Sniffing
Rat
Increased
5
7
310
Catalepsy
Rat
Increased
5
883
381
MIF-1 analog YPLG
Catalepsy
Mouse
No change
8
1051
380
MSH
Defecation
Rat
Increased
1
147
119
MSH MSH MSH
Defecation Grooming Vocalization
Rat Rat Chicken
Increased Increased Decreased
2 2 11
1S 143 333 647
120 401 413
MSH (␣)
Anxiety behavior
Rat
Increased
17
171
207
Nucleus tractus solitarius admin. Third ventricle admin. Nucleus tractus solitarius admin. ICV
ICV Foot-shock-induced behavior Paraventricular nucleus admin.
ICV Intraspinal admin. Admin. simultaneously with neuromedin B-10 Intraspinal admin. Morphine-withdrawalinduced jumping Admin. to medial preoptic area Haloperidol-induced catalepsy Haloperidol-induced catalepsy ICV; dopamine- or 5HTstimulated circling Novelty-induced defecation Striatally-evoked head turning 5HT-stimulated twitch 6-hydroxydopaminelesioned rats treated with desipramine Apomorphine- or L-DOPAinduced rotation in 6hydroxydopamine-lesioned rats Apomorphine-induced sniffing Haloperidol-induced catalepsy Haloperidol-induced catalepsy Passive-avoidance-induced defecation Novelty-induced defecation ICV Separation-induced vocalization in chicks Admin. to medial preoptic area
(continued on next page)
2230
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 7 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
MSH (␣) MSH (␣) MSH (␣)
Grooming Grooming Grooming
Rat Rat Rat
Increased Increased Increased
2 4 16
101 721 821
402 286 110
MSH (␣)
Grooming
Rat
Increased
18
393
470
MSH (␣)
Grooming
Rat
Increased
7
1
134
MSH (␣)
Grooming
Rat
Increased
Not antagonized by ␥-MSH ICV ICV; antagonized by diazepam or baclofen ICV; antagonized by melanin-concentrating hormone Enhanced by ACTH frag. (1–4) Ventral tegmental area admin.; effect enhanced by bicuculline, blocked by atropine
12
203
127
MSH (␥) MSH analog [Nle4, DPhe7]-␣ MSH MSH analogs MSH antagonist [D-Trp7, Ala8, D-Phe10) ␣-MSH(6–11)amide Neuromedin B Neuromedin B-10
Grooming Grooming
Rat Rat
No change Increased
2 4
101 721
402 286
Grooming Walking pattern
Rat Rat
Increased Increased recovery
6 16
1185 319
505 433
Bite/scratch behavior Licking
Mouse Rat
Increased Decreased
7 17
835 107
47 175
NP-E-I NPY NPY
Grooming Anxiety behavior Anxiety behaviors
Rat Rat Mouse
18 14 19
393 909 359
470 428 387
NPY NPY NPY antibody
Catalepsy Grasping Barrel rolling
Rat Rat Mouse
Increased Decreased Decreased at low dose, increased at high dose Increased Increased Increased
15 15 15
799 799 607
54 54 579
NPY antibody
Circling
Mouse
Increased
15
607
579
NPY frag. (1–30) NPY frag. (1–30) NT NT NT NT
Catalepsy Grasping Grooming Respiration Respiration Righting
Rat Rat Rat Monkey Rat Rat
No effect No effect No change Decreased No change Impaired
15 15 4 5 4 2
799 799 493 125 277 1S 99
54 54 511 366 395 187
NT NT NT
Rotation Vomiting Yawning
Rat Dog Rat
Increased Increased Decreased
6 6 12
1057 1S 173 755
389 597 398
NT analog [D-Trp11]NT
Yawning
Rat
Decreased
12
755
398
OT OT OT
Barrel rolling Grooming Grooming
Rat Rat Rat
Increased Increased Increased
6 12 9
747 113 1S 223
492 405 522
OT OT
Grooming Grooming and scratching
Rat Rat
Increased
18 2
1389 1
153 350
ICV
Admin. after sciatic nerve crush Intraspinal admin. Admin. simultaneously with gastrin-releasing peptide ICV; antagonized by MSH ICV ICV; anxiolytic effect via Y1 receptors and anxiogenic via Y2 receptors ICV ICV With admin. to ventrolateral thalamus but not other brain areas With admin. to ventromedial thalamus but not other brain areas ICV ICV PVN and IV admin. ICV ICV ICV; alcohol-induced righting impairment Substantia nigra admin. ICV ICV; apomorphine-induced yawning ICV; apomorphine-induced yawning ICV ICV Ventral tegmental area admin.; effect not altered by sex, sex-steroids or dopamine antagonists ICV With central, but not peripheral, admin.; in nonstressful, but not stressful, conditions
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2231
Table 7 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
OT
Human
Decreased
In a single patient
13
1083
139
Rat Rat Dog Rat Rat Mouse Rat
Increased Increased Increased Increased No change Increased Increased
Intrathecal admin. ICV ICV ICV Intraspinal admin. ICV
6 6 6 10 20 7 9
567 1191 1S 173 559 1445 835 475
284 538 597 11 14 47 58
Rat
Increased
ICV
9
475
58
Sauvagine
Obsessive compulsive behavior Startle response Tail movement Vomiting Yawning Anxiety behavior Bite/scratch behavior Amphetamine-induced stereotypy Apomorphine-induced stereotypy Grooming
Rat
Increased
6
3S 53
392
Sauvagine Secretin Secretin Secretin Somatostatin Somatostatin
Grooming Defecation Gnawing Grooming Barrel rolling Scratching
Rat Rat Rat Rat Rat Mouse
Decreased Increased No change No change Increased Increased
6 4 4 4 10 9
3S 53 739 739 739 1153 909
392 83 83 83 569 567
Somatostatin frag. (3–8), (9–14), or (7–10) SP SP SP
Barrel rolling
Rat
No change
ICV but not SC; in fasted, but not satiated animals SC but not ICV ICV ICV ICV ICV Intrathecal admin.; not antagonized by D-Pro2,DTrp7,9-SP and phenoxybenzamine ICV
10
1153
569
Bite/scratch behavior Biting Boxing behavior
Mouse Mouse Rat
Increased Increased Increased
Intraspinal admin. Spinal subarachnoid admin. Medial frontal cortex admin.; carbachol-induced boxing behavior
7 20 6
835 301 891
47 469 107
SP SP
Chafing movements Grooming
Fish Mouse
Increased Increased
6 6
139 363
86 217
SP SP
Grooming Grooming
Rat Rat
Increased Increased
1 7
103 557
262 352
SP SP SP SP
Licking Respiration Rigidity Risk assessment behavior
Mouse Rat Rat Rat
Increased No change Reversed Increased
20 4 1 20
301 277 103 1437
469 395 262 126
SP
Rotation
Rat
Normalized
18
1161
396
SP SP
Scratching Scratching
Mouse Mouse
Increased Increased
4 6
517 363
249 217
SP
Scratching
Mouse
Increased
9
909
567
SP SP SP SP antagonist [D-Pro2, D-Phe7, D-Trp9]-SP
Scratching Scratching and licking Vomiting Boxing behavior
Mouse Rat Dog Rat
Increased Increased No change Decreased
20 3 6 9
301 49 1S 173 1S 117
469 485 597 521
OT OT OT OT Pancreatic polypeptide Ranatensin Saralasin Saralasin
ICV; different effectiveness in different strains and ages; enhanced by naloxone but not phenoxybenzamine or haloperidol ICV ICV; not antagonized by morphine Spinal subarachnoid admin. ICV ICV; in haloperidol treated Dorsal periaqueductal gray matter admin. Animals pre-treated with peptide prior to nigrostriatal lesions; dopaminedepletion-induced turning Intrathecal admin. ICV; different effectiveness in different strains and ages Intrathecal admin.; antagonized by D-Pro2,DTrp7,9-SP and phenoxybenzamine Spinal subarachnoid admin. Intrathecal admin. ICV Carbachol-induced boxing behavior
(continued on next page)
2232
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 7 (continued) Peptide 2
Behavior
Species
Effect
Notes
Vol
Page
Ref
Carbachol-induced boxing behavior Carbachol-induced boxing behavior
9
1S 117
521
9
1S 117
521
4 20
763 1437
219 126
4
763
219
SP antagonist [D-Pro , D-Trp7,9]-SP SP antisera
Boxing behavior
Rat
Decreased
Boxing behavior
Rat
No change
SP frag. (1–7) SP frag. (7–11)
Grooming Risk assessment behavior
Mouse Rat
Decreased Increased
SP frag. analog pyroglutamyl-SP (7–11) Substance K Tachykinin agonist GR73632 Tachykinin agonist GR73632 Tachykinin agonist GR73632 Tachykinin antagonist [Sar9,Met(O2)11]SP TK PG-KII TRH
Grooming
Mouse
Increased
Bite/scratch behavior Biting
Mouse Mouse
Increased Increased
Intraspinal admin. Spinal subarachnoid admin.
7 20
835 301
47 469
Licking
Mouse
Increased
Spinal subarachnoid admin.
20
301
469
Scratching
Mouse
Increased
Spinal subarachnoid admin.
20
301
469
Grooming
Rat
Increased
18
1349
87
Grooming Grooming
Rat Rat
Increased Increased
18 9
825 283
437 565
TRH
Licking
Rat
Increased
2
1S 99
187
TRH
Respiration
Rat
Increased
4
277
395
TRH
Righting
Rat
Improved
2
1S 99
187
TRH TRH TRH analog RGH 2202 TRH analog RX 77386 TRH analog RX 77386 Urotensin I
Shaking Tail movement Grooming Grooming Shaking Grooming
Rat Rat Rat Rat Rat Rat
Increased Increased Increased Increased Increased Increased
Nucleus basalis magnocellularis admin. ICV ICV; effect antagonized by haloperidol, naloxone and NT ICV; punished licking in water-deprived, ethanoltreated rats ICV; decreased tidal volume ICV; alcohol-induced righting impairment ICV Intrathecal admin.
4 6 12 11 11 6
239 1191 1309 897 897 3S 53
155 538 152 258 258 392
Urotensin I Vasotocin VIP antagonists
Grooming Development behaviors Development behaviors
Rat Cat Mouse
Decreased Delayed Decreased
6 5 18
3S 53 25 1131
392 204 596
VP VP VP
Barrel rolling Flank marking Flank marking
Rat Hamster Hamster
Increased Increased Increased
7 14 17
213 1049 1359
290 244 5
VP
Flank marking
Hamster
Increased
16
269
4
VP VP
Grooming Grooming and scratching
Rat Rat
Increased Increased
18 2
1389 1
153 350
VP VP
Hind limb movement Scratching
Rat Gerbil
Decreased Increased
9 4
1335 801
324 311
Dorsal periaqueductal gray matter admin.
Neonates Neonates ICV but not SC; in fasted, but not satiated, animals SC but not ICV Neonates Admin. during embryogenesis; neonates ICV Medial preoptic area admin. Medial preoptic-anterior hypothalamus admin.; estradiol- and progesteronesensitive response Medial preoptic-anterior hypothalamus, central gray or lateral septum-bed nucleus admin.; testosterone-sensitive effect ICV With central, but not peripheral, admin.; in nonstressful, but not stressful, conditions Intrathecal admin. With IV, but not SC, admin.; spontaneously seizing gerbil
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2233
Table 7 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
VP
Scratching
Rat
Increased
13
17
547
VP VP and OT Z-L-glutamyl-L-leucine
Vocalization Grooming Jumping
Sheep Rat Mouse
Decreased Increased Delayed
9 15 4
1221 229 417
417 52 295
Z-L-glycyl-L-proline
Jumping
Mouse
Delayed
4
417
295
Z-L-leucyl-L-glycine
Jumping
Mouse
No change
4
417
295
Z-L-prolyl-L-leucine
Jumping
Mouse
No change
4
417
295
Z-L-prolyl-D-leucine
Jumping
Mouse
Delayed
Intrathecal admin.; potentiated by naltrexone ICV Prenatal exposure Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping
4
417
295
2234
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 8 Stereotyped and other behaviors sorted by behavior Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
AT II
Amphetamine-induced stereotypy Amphetamine-induced stereotypy
Rat
Increased
ICV
9
475
58
Rat
Decreased
2
9
285
Rat
Increased
9
475
58
CCK CCK analog Boc-CCK-4
Amphetamine-induced stereotypy Anxiety behavior Anxiety behavior
Effect antagonized by 2-bromo-ergocryptine and naloxone ICV
Rat Rat
Increased No change
16 19
1313 27
263 247
CCK frag. CCK-8
Anxiety behavior
Rat
No change
19
27
247
Galanin MCH
Anxiety behavior Anxiety behavior
Mouse Rat
No change Decreased
16 17
1283 171
556 207
MSH (␣)
Anxiety behavior
Rat
Increased
17
171
207
NPY Pancreatic polypeptide ACTH frag. (1–24) ACTH frag. (4–10) NPY
Anxiety Anxiety Anxiety Anxiety Anxiety
Rat Rat Chicken Chicken Mouse
Decreased No change Increased Increased Decreased at low dose, increased at high dose
909 1445 915 915 359
428 14 414 414 387
AT II
Apomorphine-induced stereotypy Apomorphine-induced stereotypy Apomorphine-induced stereotypy Barrel rolling Barrel rolling
Rat
Increased
ICV; anxiolytic effect via Y1 receptors and anxiogenic via Y2 receptors ICV
14 20 11 11 19
9
475
58
Rat
Increased
ICV
9
475
58
Rat
Decreased
8
39
26
Rat Mouse
Increased Increased
8 15
1089 607
201 579
OT Somatostatin Somatostatin frag. (3–8), (9–14), or (7–10) VP BBS Gastrin-releasing peptide Kassinin Neuromedin B Ranatensin SP Substance K SP
Barrel rolling Barrel rolling Barrel rolling
Rat Rat Rat
Increased Increased No change
Rats treated chronically with haloperidol ICV or IV With admin. to ventrolateral thalamus but not other brain areas ICV ICV ICV
6 10 10
747 1153 1153
492 569 569
Barrel rolling Bite/scratch behavior Bite/scratch behavior Bite/scratch behavior Bite/scratch behavior Bite/scratch behavior Bite/scratch behavior Bite/scratch behavior Biting
Rat Mouse Mouse Mouse Mouse Mouse Mouse Mouse Mouse
Increased Increased Increased Increased Increased Increased Increased Increased Increased
7 7 7 7 7 7 7 7 20
213 835 835 835 835 835 835 835 301
290 47 47 47 47 47 47 47 469
Tachykinin agonist GR73632 CRF
Biting
Mouse
Increased
20
301
469
Boxing behavior
Rat
Decreased
6
891
107
SP
Boxing behavior
Rat
Increased
6
891
107
Endorphin (␥)
Sarlasin
Saralasin Cyclo (Leu-Gly) Dynorphin frag. (1–13) NPY antibody
behavior behavior behaviors behaviors behaviors
Central nucleus of the amygdala admin. Central nucleus of the amygdala admin. ICV Admin. to medial preoptic area Admin. to medial preoptic area ICV ICV
ICV Intraspinal admin. Intraspinal admin. Intraspinal admin. Intraspinal admin. Intraspinal admin. Intraspinal admin. Intraspinal admin. Spinal subarachnoid admin. Spinal subarachnoid admin. Medial frontal cortex admin.; carbacholinduced boxing behavior Medial frontal cortex admin.; carbacholinduced boxing behavior
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2235
Table 8 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
SP antagonist [D-Pro , D-Phe7, D-Trp9]-SP SP antagonist [D-Pro2, D-Trp7,9]-SP SP antisera
Boxing behavior
Rat
Decreased
9
1S 117
521
Boxing behavior
Rat
Decreased
9
1S 117
521
Boxing behavior
Rat
No change
9
1S 117
521
CCK-8
Catalepsy
Rat
Blocked
6
237
253
Ceruletide
Catalepsy
Rat
Decreased
10
779
250
Ceruletide analogs
Catalepsy
Mouse
Increased
3
701
610
Dermorphin
Catalepsy
Rat
Increased
6
3S 165
59
Dermorphin
Catalepsy
Rat
Decreased
6
3S 165
59
Endorphin ()
Catalepsy
Rat
Increased
6
237
253
MIF-1
Catalepsy
Mouse
Prevented tolerance
8
1051
380
MIF-1
Catalepsy
Rat
Decreased
2
105
85
MIF-1 analog parepide
Catalepsy
Rat
Increased
5
883
381
MIF-1 analog YPLG
Catalepsy
Mouse
No change
8
1051
380
NPY NPY frag. (1–30) SP CCK-4 CCK-8 CCK-8, desulfated Dynorphin frag. (1–6) Dynorphin frag. (1–7) Dynorphin frag. (1–8)
Catalepsy Catalepsy Chafing movements Circling Circling Circling Circling Circling Circling
Rat Rat Fish Rat Rat Rat Rat Rat Rat
Increased No effect Increased No change Increased No change Increased Increased Increased
Carbachol-induced boxing behavior Carbachol-induced boxing behavior Carbachol-induced boxing behavior ICV; endorphininduced catalepsy; effect blocked by frontal cortex lesion Haloperidol-induced catalepsy Hexobarbital-induced sleep ICV; effects increased by caudate lesion ICV; in dermorphintolerant, but not morphine-tolerant, rats ICV; effects enhanced by frontal cortex ablation Haloperidol-induced catalepsy Haloperidol-induced catalepsy Haloperidol-induced catalepsy Haloperidol-induced catalepsy ICV ICV
15 15 6 7 7 7 8 8 8
799 799 139 809 809 809 837 837 837
54 54 86 367 367 367 186 186 186
Dynorphin frag. (2–17) Enkephalin (Leu) MIF-1
Circling Circling Circling
Rat Rat Rat
Increased Increased No change
8 8 2
837 837 189
186 186 148
NPY antibody
Circling
Mouse
Increased
15
607
579
Ceruletide analogs
Convulsions
Mouse
Decreased
3
701
610
CCK-4
Defecation
Rat
No change
6
91
275
CCK-8
Defecation
Rat
No change
6
91
275
MIF-1
Defecation
Rat
Decreased
2
1S 143
120
MSH
Defecation
Rat
Increased
1
147
119
2
ICV ICV ICV Substantia nigra admin. Substantia nigra admin. Substantia nigra admin.; not antagonized by naloxone Substantia nigra admin. Substantia nigra admin. ICV; dopamine- or 5HT-stimulated circling With admin. to ventromedial thalamus, but not other brain areas Harman-induced convulsions Nucleus accumbens admin. Nucleus accumbens admin. Novelty-induced defecation Passive-avoidanceinduced defecation
(continued on next page)
2236
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 8 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
MSH
Defecation
Rat
Increased
2
1S 143
120
Secretin MIF-1 Vasotocin VIP antagonists
Defecation Depressive behaviors Development behaviors Development behaviors
Rat Human Cat Mouse
Increased Decreased Delayed Decreased
Novelty-induced defecation ICV
4 4 5 18
739 297 25 1131
83 559 204 596
ACTH ACTH frag. (4–10) Casomorphin frag. (1–5) Endorphin (␣)
Distress Distress Distress Distress
vocalization vocalization vocalization vocalization
Chicken Chicken Chicken Chicken
Increased Increased Decreased Decreased
11 11 5 5
915 915 829 823
414 414 415 573
Endorphin ()
Distress vocalization
Chicken
Decreased
5
823
573
Endorphin (␥)
Distress vocalization
Chicken
Decreased
5
823
573
VP
Flank marking
Hamster
Increased
14
1049
244
VP
Flank marking
Hamster
Increased
17
1359
5
VP
Flank marking
Hamster
Increased
16
269
4
CRF antagonist ␣-helical CRF Galanin
Freezing behavior
Rat
Decreased
15
1303
237
Freezing behavior
Rat
No change
15
1303
237
Ceruletide analogs
Gnawing
Mouse
Increased
2
2S 65
609
Secretin NPY NPY frag. (1–30) ACTH
Gnawing Grasping Grasping Grooming
Rat Rat Rat Rat
No change Increased No effect Increased
4 15 15 7
739 799 799 597
83 54 54 476
ACTH ACTH
Grooming Grooming
Rat Rat
Increased Increased
4 5
907 713
202 546
ACTH ACTH
Grooming Grooming
Rat Rat
Increased Increased
6 7
369 651
562 160
ACTH
Grooming
Rat
Increased
16
1263
434
ACTH
Grooming
Rat and mouse
Increased
8
841
159
ACTH frag. (1–24)
Grooming
Rat
Increased
4
833
6
ACTH frag. (1–4)
Grooming
Rat
Enhanced
7
1
134
ACTH/MSH frag. analogs, C-frag. elongated or cyclized
Grooming
Rat
Increased
5
1197
232
Neonates Admin. during embryogenesis; neonates
ICV; chicks With ICV, but not IP, admin. With ICV, but not IP, admin.; chicks With ICV, but not IP, admin. Medial preoptic area admin. Medial preopticanterior hypothalamus admin.; estradiol- and progesterone-sensitive response Medial preopticanterior hypothalamus, central gray or lateral septum-bed nucleus admin.; testosteronesensitive effect Foot-shock-induced behavior Foot-shock-induced behavior Methylphen-induced gnawing ICV ICV ICV ICV; blocked by naloxone ICV Pre-optic, anterior hypothalamic admin. ICV ICV; area of action localized to anteriorventral third ventricle ICV; not antagonized by nitric oxide synthase inhibitor ICV; greater effect in rats than mice ICV; effect antagonized by naloxone In MSH-induced grooming; no effect alone More potent than unmodified frag.
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2237
Table 8 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
Amylin BBS BBS
Grooming Grooming Grooming
Rat Rat Rat
No change Increased Increased
17 2 8
589 2S 179 237
91 198 303
BBS BBS
Grooming Grooming
Rat Rat
Increased Increased
12 13
761 1215
182 99
BBS
Grooming
Rat
Increased
4
693
355
BBS
Grooming
Rat
Increased
7
557
352
BBS
Grooming
Rat
Increased
9
1S 233
260
BBS
Grooming
Rat
Increased
9
1S 245
260
BBS
Grooming
Rat
Increased
4
907
202
BBS
Grooming
Rat
Increased
6
369
562
BBS
Grooming
Rat
Increased
6
1179
564
BBS
Grooming
Rat
Increased
6
1179
563
CGRP
Grooming
Rat
Increased
17
1183
294
CGRP CRF CRF CRF
Grooming Grooming Grooming Grooming
Rat Rat Rat Rat
Decreased Increased Increased Increased
5 13 6 20
861 1149 3S 53 509
297 146 392 571
CRF
Grooming
Rat and mouse
Increased
8
841
159
Dynorphin analog desTyr1-dynorphin (1–13) Dynorphin frag. (1–13)
Grooming
Rat
Increased
4
833
6
Grooming
Rat
Increased
4
833
6
Dynorphin frag. (1–13)
Grooming
Rat and mouse
Increased
1
341
578
Galanin
Grooming
Rat
No change
11
1001
304
MSH MSH (␣)
Grooming Grooming
Rat Rat
Increased Increased
2 2
333 101
401 402
MSH (␣) MSH (␣)
Grooming Grooming
Rat Rat
Increased Increased
ICV ICV With admin. to multiple brain areas ICV ICV; developed over 8 days ICV; effect antagonized by neuroleptics ICV; antagonized by morphine With admin. to nucleus tractus solitarius but not nucleus accumbens With admin. to nucleus tractus solitarius but not nucleus accumbens With ICV, but not IV, admin.; no antagonism by haloperidol, morphine, naloxone, or NT. No effect of hypophysectomy or adrenalectomy ICV; different doses induce different types of grooming Effect enhanced by haloperidol, antagonized by naloxone and NT ICV; antagonized by haloperidol Antagonized by nitric oxide synthase inhibitor ICV ICV ICV but not SC ICV; not antagonized by melacortin-4 receptor antagonist ICV; greater effect in rats than mice; not antagonized by dexamethasone ICV; effect antagonized by naloxone ICV; effect antagonized by naloxone ICV; not reversible by naloxone Paraventicular nucleus admin. ICV Not antagonized by ␥ MSH ICV ICV; antagonized by diazepam or baclofen
4 16
721 821
286 110
(continued on next page)
2238
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 8 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
MSH (␣)
Grooming
Rat
Increased
18
393
470
MSH (␣)
Grooming
Rat
Increased
7
1
134
MSH (␣)
Grooming
Rat
Increased
ICV; antagonized by melanin-concentrating hormone Enhanced by ACTH frag. (1–4) Ventral tegmental area admin.; effect enhanced by bicuculline, blocked by atropine
12
203
127
MSH (␥) MSH analog [Nle4, D-Phe7]-␣ MSH MSH analogs NP-E-I
Grooming Grooming
Rat Rat
No change Increased
2 4
101 721
402 286
Grooming Grooming
Rat Rat
Increased Increased
6 18
1185 393
505 470
NT OT OT
Grooming Grooming Grooming
Rat Rat Rat
No change Increased Increased
4 12 9
493 113 1s 223
511 405 522
OT Sauvagine
Grooming Grooming
Rat Rat
Increased Increased
18 6
1389 3S 53
153 392
Sauvagine Secretin SP
Grooming Grooming Grooming
Rat Rat Mouse
Decreased No change Increased
6 4 6
3S 53 739 363
392 83 217
SP SP
Grooming Grooming
Rat Rat
Increased Increased
1 7
103 557
262 352
SP frag. (1–7) SP frag. analog pyroglutamyl-SP (7–11) Tachykinin antagonist [Sar9,Met(O2)11]SP TK PG-KII TRH
Grooming Grooming
Mouse Mouse
Decreased Increased
4 4
763 763
219 219
Grooming
Rat
Increased
18
1349
87
Grooming Grooming
Rat Rat
Increased Increased
18 9
825 283
437 565
TRH analog RGH 2202 TRH analog RX 77386 Urotensin I
Grooming Grooming Grooming
Rat Rat Rat
Increased Increased Increased
12 11 6
1309 897 3S 53
152 258 392
Urotensin I VP VP and OT BBS
Grooming Grooming Grooming Grooming and scratching
Rat Rat Rat Rat
Decreased Increased Increased Increased
6 18 15 10
3S 53 1389 229 529
392 153 52 259
ICV
ICV; antagonized by MSH PVN and IV admin. ICV Ventral tegmental area admin.; effect not altered by sex, sexsteroids, or dopamine antagonists ICV ICV but not SC; in fasted, but not satiated, animals SC but not ICV ICV ICV; different effectiveness in different strains and ages; enhanced by naloxone but not phenoxybenzamine or haloperidol ICV ICV; not antagonized by morphine
Nucleus basalis magnocellularis admin. ICV ICV; effect antagonized by haloperidol, naloxone and NT Neonates ICV but not SC; in fasted, but not satiated, animals SC but not ICV ICV Prenatal exposure Neonates
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2239
Table 8 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
OT
Grooming and scratching
Rat
Increased
2
1
350
VP
Grooming and scratching
Rat
Increased
2
1
350
MIF-1
Head turning
Rat
No change
2
189
148
MIF-1 VP Dynorphin frag. (1–13) CRF
Mouse Rat Rat Monkey
No change Decreased Increased Increased
2 9 4 4
189 1335 631 217
148 324 168 268
Casomorphin analog BCH 325 L-pyroglutamyl-Lleucine Z-L-glutamyl-L-leucine
Head twitch Hind limb movement Hind limb paralysis Huddling and lying down Jerking
With central, but not peripheral, admin.; in non-stressful, but not stressful, conditions With central, but not peripheral, admin.; in non-stressful, but not stressful, conditions Striatally-evoked head turning 5HT-stimulated twitch Intrathecal admin. Intrathecal admin. ICV
Rat
Decreased
16
635
447
Jumping
Mouse
Delayed
4
417
295
Jumping
Mouse
Delayed
4
417
295
Z-L-glycyl-L-proline
Jumping
Mouse
Delayed
4
417
295
Z-L-leucyl-L-glycine
Jumping
Mouse
No change
4
417
295
Z-L-prolyl-L-leucine
Jumping
Mouse
No change
4
417
295
Z-L-prolyl-D-leucine
Jumping
Mouse
Delayed
Bromocriptine-induced jerks Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping
4
417
295
BBS BBS Gastrin-releasing peptide
Licking Licking Licking
Rat Rat Rat
Decreased Decreased Decreased
17 16 17
107 903 107
175 523 175
Gastrin-releasing peptide Neuromedin B-10
Licking Licking
Rat Rat
Decreased Decreased
16 17
903 107
523 175
SP
Licking
Mouse
Increased
20
301
469
Tachykinin agonist GR73632 TRH
Licking
Mouse
Increased
20
301
469
Licking
Rat
Increased
2
1S 99
187
Morphine-induced behaviors Myoclonus (serotonergic behavior) Obsessive compulsive behavior Opiate withdrawal behavior Oral act. (serotonergic behavior)
Mouse
No change
17
1291
97
Mouse
No change
10
5
439
Human
Decreased
13
1083
139
Rat
No change
15
1303
237
Rat
No change
14
1159
206
Mouse Rat Rat
Increased Increased Increased
2 4 4
2S 65 277 277
609 395 395
Adrenal Peptide E (Frog) ACTH OT Galanin MIF-1
Ceruletide analogs BBS Calcitonin
Ptosis Respiration Respiration
Admin. simultaneously with neuromedin B-10 Admin. simultaneously with gastrin-releasing peptide Spinal subarachnoid admin. Spinal subarachnoid admin. ICV; punished licking in water-deprived, ethanol-treated rats ICV 5-hydroxytryptophaninduced In a single patient
6-hydroxydopaminelesioned rats treated with desipramine ICV ICV
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Table 8 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
CCK Enkephalin analog DADLE Enkephalin analog DADLE Enkephalin analog DAGO Enkephalin analog FK33–824 NT NT SP TRH
Respiration Respiration
Cat Rat
Increased tidal volume Increased
9 3
809 1031
246 225
Respiration
Rat
Increased
Ventral medulla admin. Nucleus tractus solitarius admin. Third ventricle admin.
3
1023
236
Respiration
Rat
Nucleus tractus solitarius admin.
3
1031
225
Respiration
Mouse
Increased at low dose decreased at high dose Decreased rate
9
777
378
Respiration Respiration Respiration Respiration
Monkey Rat Rat Rat
Decreased No change No change Increased
5 4 4 4
125 277 277 277
366 395 395 395
BBS
Righting
Rat
Impaired
2
1S 99
187
Endorphin ()
Righting
Rat
Impaired
2
1S 99
187
NT
Righting
Rat
Impaired
2
1S 99
187
TRH
Righting
Rat
Improved
2
1S 99
187
SP
Rigidity
Rat
Reversed
1
103
262
SP
Risk assessment behavior
Rat
Increased
20
1437
126
SP frag. (7–11)
Risk assessment behavior
Rat
Increased
20
1437
126
ACTH/MSH 4–10 analog ORG 2766
Rotation
Rat
Decreased
14
1317
10
CCK-8
Rotation
Rat
Increased
18
1161
396
Cyclo (Leu-Gly)
Rotation
Rat
No change
8
39
26
MIF-1 analog 3(R)-[(2(S)pyrrolidinylcarbonyl) amino]-2-oxo-1pyrrolidineacetamide NT SP
Rotation
Rat
Increased
ICV ICV ICV ICV; decreased tidal volume ICV; alcohol-induced righting impairment ICV; alcohol-induced righting impairment ICV; alcohol-induced righting impairment ICV; alcohol-induced righting impairment ICV; in haloperidol treated Dorsal periaqueductal gray matter admin. Dorsal periaqueductal gray matter admin. Nigrostriatal lesions and amphetamineinduced turning Animals pre-treated with peptide prior to nigrostriatal lesions; dopamine-depletioninduced turning Apomorphine-induced rotation in rats with unilateral nigrostriatal lesions Apomorphine- or LDOPA-induced rotation in 6-hydroxydopaminelesioned rats
18
1209
363
Rotation Rotation
Rat Rat
Increased Normalized
6 18
1057 1161
389 396
CCK-8 Gastrin (1–17) BBS
Rumination Rumination Scratching
Sheep Sheep Mouse
No change Increased Increased
5 5 9
81 81 909
238 238 567
BBS
Scratching
Rat
Increased
6
1179
563
Substantia nigra admin. Animals pre-treated with peptide prior to nigrostriatal lesions; dopamine-depletioninduced turning ICV Intrathecal admin.; not antagonized by DPro2,D-Trp7,9-SP and phenoxybenzamine ICV; antagonized by naloxone
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2241
Table 8 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
Somatostatin
Scratching
Mouse
Increased
9
909
567
SP SP
Scratching Scratching
Mouse Mouse
Increased Increased
4 6
517 363
249 217
SP
Scratching
Mouse
Increased
9
909
567
SP
Scratching
Mouse
Increased
20
301
469
Tachykinin agonist GR73632 VP
Scratching
Mouse
Increased
20
301
469
Scratching
Gerbil
Increased
4
801
311
VP
Scratching
Rat
Increased
13
17
547
SP Dynorphin frag. (1–13) Enkephalin (Met) Enkephalin analog DAMA TRH TRH analog RX 77386 BBS MIF-1 analog Cyclo(Leu-Gly) Endorphin () Melatonin OT CCK
Scratching and licking Shaking Shaking Shaking
Rat Rat Fish Rat
Increased Increased Increased Increased
Intrathecal admin.; not antagonized by DPro2,D-Trp7,9-SP and phenoxybenzamine Intrathecal admin. ICV; different effectiveness in different strains and ages Intrathecal admin.; antagonized by DPro2,D-Trp7,9-SP and phenoxybenzamine Spinal subarachnoid admin. Spinal subarachnoid admin. With IV, but not SC, admin.; spontaneously seizing gerbil Intrathecal admin.; potentiated by naltrexone Intrathecal admin. ICV or IV
3 8 6 4
49 1089 139 239
485 201 86 155
Shaking Shaking Sighing Sniffing
Rat Rat Rat Rat
Increased Increased Increased Increased
4 11 4 5
239 897 277 7
155 258 395 310
Startle Startle response Startle response Stretching
Rat Rat Rat Mouse
Decreased Decreased Increased Decreased
2 2 6 10
1S 137 1S 155 567 31
234 188 284 9
OT TRH Endorphin ()
Tail movement Tail movement Trembling
Rat Rat Rat
Increased Increased Decreased
6 6 2
1191 1191 1S 99
538 538 187
Enkephalin MSH
Vocalization Vocalization
Sheep Chicken
Decreased Decreased
9 11
1221 647
417 413
VP ACTH AT BBS Bradykinin NT OT SP MSH antagonist [D-Trp7, Ala8, DPhe10)␣-MSH (6–11)amide Calcitonin
Vocalization Vomiting Vomiting Vomiting Vomiting Vomiting Vomiting Vomiting Walking pattern
Sheep Dog Dog Dog Dog Dog Dog Dog Rat
Decreased No change No change No change No change Increased Increased No change Increased recovery
9 6 6 6 6 6 6 6 16
1221 1S 173 1S 173 1S 173 1S 173 1S 173 1S 173 1S 173 319
417 597 597 597 597 597 597 597 433
Writhing
Mouse
Increased analgesia
6
3S 277
200
ICV ICV Neonates ICV Apomorphine-induced Sniffing
ICV; abdominal irritant-induced stretching Intrathecal admin. Intrathecal admin. ICV; alcohol-induced tremors ICV Separation-induced vocalization in chicks ICV ICV ICV ICV ICV ICV ICV ICV Admin. after sciatic nerve crush
ICV, IP, and IV, but no effect SC; Achinduced writhing
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Table 8 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
ACTH
Yawning
Rat
Increased
16
1263
434
Casomorphin analog BCH 325
Yawning
Rat
Normalized
16
635
447
Casomorphin analogs
Yawning
Rat
Decreased
13
69
464
NT
Yawning
Rat
Decreased
12
755
398
NT analog[D-Trp11]NT
Yawning
Rat
Decreased
12
755
398
OT
Yawning
Rat
Increased
ICV; antagonized by nitric oxide synthase inhibitor Bromocriptine-evoked yawning that was inhibited by reserpine ICV; apomorphineinduced yawning ICV; apomorphineinduced yawning ICV; apomorphineinduced yawning ICV
10
559
11
2.7. Stereotyped and other behaviors No categorization system is likely to capture all of the types of behaviors that peptides can affect. Because readers might be interested in a particular behavior listed in this catch-all category, the table is presented twice, once organized by peptide (Table 7) and once by behavior (Table 8).
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R.N. McLay et al. / Peptides 22 (2001) 2181–2255 [117] Danguir J. Food intake in rats is increased by intracerebroventricular infusion of the somatostatin analogue SMS 201–995 and is decreased by somatostatin antiserum. Peptides 1988;9:211–3. [118] Datta PC, Hoehler FK, Sandman CA. Effects of melatonin on startle reflex in rat. Peptides 1981;2 Suppl 1:155– 60. [119] Datta PC, King MG. Effects of ␣-MSH and melatonin on passive avoidance and on PA-induced defecation and plasma 11-OHCS in hypophysectomized rats. Peptides 1980;1:147–53. [120] Datta PC, King MG. ␣-MSH, MIF-I, and melatonin: effects on novelty-induced defecation, plasma 11-OHCS, and central catecholamines in rats. Peptides 1981;2 Suppl 1:143–54. [121] Datta PC, Sandman CA, Hoehler FK. Attenuation of morphine analgesia by ␣-MSH, MIF-I, melatonin and naloxone in the rat. Peptides 1982;3:433–7. [122] Davis JL, Pico RM. Arginine vasotocin delays extinction of a conditioned avoidance behavior in neonatal chicks. Peptides 1984; 5:1221–3. [123] Davis JL, Pico RM, Cherkin A. L-prolyl-L-arginyl-glycinamide induces memory enhancement in chicks. Peptides 1983;4:401– 4. [124] Davis JM, Lowy MT, Yim GK, Lamb DR, Malven PV. Relationship between plasma concentrations of immunoreactive -endorphin and food intake in rats. Peptides 1983;4:79 – 83. [125] Davis TP, Gillespie TJ, Porreca F. Peptide fragments derived from the -chain of hemoglobin (hemorphins) are centrally active in vivo. Peptides 1989;10:747–51. [126] De Araujo JE, Silva RC, Huston JP, Brandao ML. Anxiogenic effects of substance P and its 7–11 C terminal, but not the 1–7 N terminal, injected into the dorsal periaqueductal gray. Peptides 1999; 20:1437– 43. [127] de Barioglio SR, Lezcano N, Celis ME. Alpha MSH-induced excessive grooming behavior involves a GABAergic mechanism. Peptides 1991;12:203–5. [128] de Beaurepaire R, Suaudeau C. Anorectic effect of calcitonin, neurotensin and bombesin infused in the area of the rostral part of the nucleus of the tractus solitarius in the rat. Peptides 1988;9:729 –33. [129] De Caro G, Massi M. Water intake modifications induced by tachykinins, bombesins and opioid peptides. Peptides 1985;6(Suppl 3): 181–5. [130] De Caro G, Massi M, Micossi LG, Perfumi M. Angiotensin II antagonists versus drinking induced by bombesin or eledoisin in pigeons. Peptides 1982;3:631– 6. [131] De Caro G, Massi M, Micossi LG, Perfumi M. Drinking and feeding inhibition by ICV pulse injection or infusion of bombesin, ranatensin and litorin to rats. Peptides 1984;5:607–13. [132] de Castiglione R. Structural requirements of ceruletide-like peptides for activity on gut and brain. Peptides 1981;2 Suppl 2:61–3. [133] De Ceballos ML, Lopez AE, Harto JR, Bravo A, Gomez-Monterrey I, Gonzalez-Muniz R, Garcia-Lopez MT, Del Rio J. Prolonged antinociceptive activity of pseudodipeptide analogues of LysTrp(Nps) and Trp(Nps)-Lys. Peptides 1992;13:63–7. [134] De Graan PN, Spruijt BM, Eberle AN, Girard J, Gispen WH. ACTH1– 4 potentiates ␣-MSH-induced melanophore dispersion, and excessive grooming. Peptides 1986;7:1– 4. [135] de Pedro N, Cespedes MV, Delgado MJ, Alonso-Bedate M. Muopioid receptor is involved in -endorphin-induced feeding in goldfish. Peptides 1996;17:421– 4. [136] de Pedro N, Pinillos ML, Valenciano AI, Alonso-Bedate M, Delgado MJ. Inhibitory effect of serotonin on feeding behavior in goldfish: involvement of CRF. Peptides 1998;19:505–11. [137] Della-Fera MA, Baile CA. CCK-octapeptide injected in CSF decreases meal size, and daily food intake in sheep. Peptides 1980;1: 51– 4. [138] Della-Fera MA, Baile CA, Beinfeld MC. Cerebral ventricular transport and uptake: importance for CCK-mediated satiety. Peptides 1982;3:963– 8. [139] den Boer JA, Westenberg HG. Oxytocin in obsessive compulsive disorder. Peptides 1992;13:1083–5.
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[140] Denbow DM. Centrally and peripherally administered bombesin decreases food intake in turkeys. Peptides 1989;10:275–9. [141] Denbow DM, Duke GE, Chaplin SB. Food intake, gastric secretion, and motility as affected by avian pancreatic polypeptide administered centrally in chickens. Peptides 1988;9:449 –54. [142] Denbow DM, Myers RD. Eating, drinking, and temperature responses to intracerebroventricular cholecystokinin in the chick. Peptides 1982;3:739 – 43. [143] Deupree D, Hsiao S. Cholecystokinin octapeptide, proglumide, and passive avoidance in rats. Peptides 1987;8:25– 8. [144] Deutch AY, Maggio JE, Bannon MJ, Kalivas PW, Tam SY, Goldstein M, Roth RH. Substance K, and substance P differentially modulate mesolimbic, and mesocortical systems. Peptides 6 Suppl 1985;2:113–22. [145] Deviche P, Schepers G. Intracerebroventricular injection of ostrich -endorphin to satiated pigeons induces hyperphagia but not hyperdipsia. Peptides 1984;5:691– 4. [146] Diamant M, Croiset G, De Wied D. The effect of corticotropinreleasing factor (CRF) on autonomic and behavioral responses during shock-prod burying test in rats. Peptides 1992;13:1149 –58. [147] Dickinson SL, Slater P. Opiate receptor antagonism by L-prolyl-Lleucyl-glycinamide, MIF-I. Peptides 1980;1:293–9. [148] Dickinson SL, Slater P. Effect of L-prolyl-L-leucyl-glycinamide (MIF-I) on some neurotransmitter-receptor interactions. Peptides 1981;2:189 –95. [149] Dickson PR, Feifel D, Vaccarino FJ. Blockade of endogenous GRF at dark onset selectively suppresses protein intake. Peptides 1995; 16:7–9. [150] Dickson PR, Vaccarino FJ. GRF-induced feeding: evidence for protein selectivity, and opiate involvement. Peptides 1994;15:1343– 52. [151] Dluzen DE, Muraoka S, Engelmann M, Landgraf R. The effects of infusion of arginine vasopressin, oxytocin, or their antagonists into the olfactory bulb upon social recognition responses in male rats. Peptides 1998;19:999 –1005. [152] Drago F, Grassi M, Valerio C, Coppi G, Lauria N, Nicotra GC, Raffaele R. Behavioral changes induced by the thyrotropin-releasing hormone analogue, RGH 2202. Peptides 1991;12:1309 –13. [153] Drago F, Stanciu M, Salehi S, Scapagnini U. The block of central vasopressin V1 but not V2 receptors suppresses grooming behavior and hypothermia induced by intracerebroventricular vasopressin in male rats. Peptides 1997;18:1389 –92. [154] Drucker-Colin R, Bernal-Pedraza J, Fernandez-Cancino F, Oksenberg A. Is vasoactive intestinal polypeptide (VIP) a sleep factor? Peptides 1984;5:837– 40. [155] Drust EG, Crawford IL. Comparison of the effects of TRH and D-Ala2-metenkephalinamide on hippocampal electrical activity and behavior in the unanesthetized rat. Peptides 1983;4:239 – 43. [156] Dryden S, Pickavance L, Henderson L, Williams G. Hyperphagia induced by hypoglycemia in rats is independent of leptin and hypothalamic neuropeptide Y (NPY). Peptides 1998;19:1549 –55. [157] Dudley CA, Vale W, Rivier J, Moss RL. The effect of LHRH antagonist analogs and an antibody to LHRH on mating behavior in female rats. Peptides 1981;2:393– 6. [158] Dulawa SC, Vanderweele DA. Cholecystokinin and estradiol synergistically potentiate satiety in rats. Peptides 1994;15:913– 8. [159] Dunn AJ, Berridge CW, Lai YI, Yachabach TL. CRF-induced excessive grooming behavior in rats, and mice. Peptides 1987;8:841– 4. [160] Dunn AJ, Hurd RW. ACTH acts via an anterior ventral third ventricular site to elicit grooming behavior. Peptides 1986;7:651–7. [161] Eaves M, Thatcher-Britton K, Rivier J, Vale W, Koob GF. Effects of corticotropin releasing factor on locomotor activity in hypophysectomized rats. Peptides 1985;6:923– 6. [162] Eckel LA, Geary N. Endogenous cholecystokinin’s satiating action increases during estrus in female rats. Peptides 1999;20:451– 6.
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R.N. McLay et al. / Peptides 22 (2001) 2181–2255 [483] Schulz S, Diedrich M, Wetzel W, Matthies H. Effects of [des-TyrD-Phe3]-casomorphin (2–5) on sleep pattern in rats. Peptides 1991; 12:893– 6. [484] Schumann MD, Payza K, Ho KK, Burgess K. FMRFamide containing E-2,3-methanomethionine, and E-2,3-methanophenylalanine. Peptides 1996;17:83– 6. [485] Seybold VS, Hylden JL, Wilcox GL. Intrathecal substance P and somatostatin in rats: behaviors indicative of sensation. Peptides 1982;3:49 –54. [486] Shaikh MB, Shaikh AB, Siegel A. Opioid peptides within the midbrain periaqueductal gray suppress affective defense behavior in the cat. Peptides 1988;9:999 –1004. [487] Shen Y, Li R. The role of neuropeptides in learning and memory: possible mechanisms. Med Hypotheses 1995;45:529 –38. [488] Shlik J, Koszycki D, Bradwejn J. Decrease in short-term memory function induced by CCK-4 in healthy volunteers. Peptides 1998; 19:969 –75. [489] Sills TL, Vaccarino FJ. Individual differences in the feeding response to CCKB antagonists: role of the nucleus accumbens. Peptides 1996;17:593–9. [490] Silver AJ, Flood JF, Morley JE. Effect of gastrointestinal peptides on ingestion in old and young mice. Peptides 1988;9:221–5. [491] Simansky KJ, Smith GP. Acute abdominal vagotomy reduces drinking to peripheral but not central angiotensin II. Peptides 1983;4: 159 – 63. [492] Skofitsch G, Jacobowitz DM. Calcitonin gene-related peptide coexists with substance P in capsaicin sensitive neurons and sensory ganglia of the rat. Peptides 1985;6:747–54. [493] Skopkova J, Croiset G, De Wied D. Differential effects of DGAVP on acquisition and extinction of active avoidance behavior. Peptides 1991;12:471–5. [494] Skopkova J, Lat J, Dvorska I, Hrbas P, Radil T. The effect of AVP and DGAVP on the exploratory activity of rats. Peptides 1987;8: 785–90. [495] Sleeman M, Willis GL, Smith GC. Cholecystokinin, behavioral tolerance, and tumor-induced anorexia. Peptides 1987;8:223– 6. [496] Smith BK, Berthoud HR, York DA, Bray GA. Differential effects of baseline macronutrient preferences on macronutrient selection after galanin, NPY, and an overnight fast. Peptides 1997;18:207–11. [497] Smith BK, York DA, Bray GA. Chronic cerebroventricular galanin does not induce sustained hyperphagia or obesity. Peptides 1994; 15:1267–72. [498] Smith GP, Jerome C, Gibbs J. Abdominal vagotomy does not block the satiety effect of bombesin in the rat. Peptides 1981;2:409 –11. [499] Smith GP, Jerome C, Kulkosky PJ, Simansky KJ. Ceruletide acts in the abdomen, not in the brain, to produce satiety. Peptides 1984;5: 1149 –57. [500] Smith J, Perez S, Rushing PA, Smith GP, Gibbs J. Gastrin-releasing peptide1–27, unlike bombesin, does not reduce sham feeding in rats. Peptides 1997;18:1465–7. [501] Smotherman WP, Levine S. ACTH4 –10 affects behavior but not plasma corticosterone levels in a conditioned taste aversion situation. Peptides 1980;1:207–10. [502] Sonti G, Ilyin SE, Plata-Salaman CR. Neuropeptide Y blocks, and reverses interleukin-1 (IL-1) -induced anorexia in rats. Peptides 1996;17:517–20. [503] South EH, Ritter RC. Capsaicin application to central or peripheral vagal fibers attenuates CCK satiety. Peptides 1988;9:601–12. [504] Sprick U, Hasenohrl RU, Krauth J, Klapdor K, Huston JP. Effects of chronic substance P treatment and i.c. fetal grafts on learning after hippocampal kainic acid lesions. Peptides 1996;17:275– 85. [505] Spruijt BM, De Graan PN, Eberle AN, Gispen WH. Comparison of structural requirements of ␣-MSH and ACTH for inducing excessive grooming and pigment dispersion. Peptides 1985;6:1185–9. [506] Stacher G, Steinringer H, Schmierer G, Schneider C, Winklehner S. Ceruletide decreases food intake in non-obese man. Peptides 1982; 3:607–12.
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[507] Stacher G, Steinringer H, Schmierer G, Schneider C, Winklehner S. Cholecystokinin octapeptide decreases intake of solid food in man. Peptides 1982;3:133– 6. [508] Stacher G, Steinringer H, Schmierer G, Winklehner S, Schneider C. Ceruletide increases threshold and tolerance to experimentally induced pain in healthy man. Peptides 1982;3:955– 62. [509] Stamidis H, Young GA. Mu-␦ opioid interactions. III: Differential antagonism of DPDPE-induced increases in morphine EEG, and EEG power spectra by DALCE, and naltrindole. Peptides 1993;14: 511–7. [510] Stanley BG, Daniel DR, Chin AS, Leibowitz SF. Paraventricular nucleus injections of peptide YY and neuropeptide Y preferentially enhance carbohydrate ingestion. Peptides 1985;6:1205–11. [511] Stanley BG, Hoebel BG, Leibowitz SF. Neurotensin: effects of hypothalamic, and intravenous injections on eating, and drinking in rats. Peptides 1983;4:493–500. [512] Stanley BG, Kyrkouli SE, Lampert S, Leibowitz SF. Neuropeptide Y chronically injected into the hypothalamus: a powerful neurochemical inducer of hyperphagia, and obesity. Peptides 1986;7: 1189 –92. [513] Stanley BG, Magdalin W, Seirafi A, Nguyen MM, Leibowitz SF. Evidence for neuropeptide Y mediation of eating produced by food deprivation and for a variant of the Y1 receptor mediating this peptide’s effect. Peptides 1992;13:581–7. [514] Stanley BG, Thomas WJ. Feeding responses to perifornical hypothalamic injection of neuropeptide Y in relation to circadian rhythms of eating behavior. Peptides 1993;14:475– 81. [515] Steiger A, Guldner J, Knisatschek H, Rothe B, Lauer C, Holsboer F. Effects of an ACTH/MSH(4 –9) analog (HOE 427) on the sleep EEG and nocturnal hormonal secretion in humans. Peptides 1991; 12:1007–10. [516] Steiger A, Holsboer F. Neuropeptides and human sleep. Sleep 2001; 20:1038 –52. [517] Stein LJ, Woods SC. Gastrin releasing peptide reduces meal size in rats. Peptides 1982;3:833–5. [518] Steinman JL, Fujikawa DG, Wasterlain CG, Cherkin A, Morley JE. The effects of adrenergic, opioid and pancreatic polypeptidergic compounds on feeding and other behaviors in neonatal leghorn chicks. Peptides 1987;8:585–92. [519] Stewart JM, Hall ME, Harkins J, Frederickson RC, Terenius L, Hokfelt T, Krivoy WA. A fragment of substance P with specific central activity: SP(1–7). Peptides 1982;3:851–7. [520] Stickrod G, Kimble DP, Smotherman WP. Met-enkephalin effects on associations formed in utero. Peptides 1982;3:881–3. [521] Stivers JA, Crawley JN. Substance P antagonists block carbacholinduced “boxing” behavior at a site of coexistence in the rat prefrontal cortex. Peptides 1988;9(Suppl 1):117–21. [522] Stivers JA, Kaltwasser MT, Hill PS, Hruby VJ, Crawley JN. Ventral tegmental oxytocin induces grooming. Peptides 1988;9(Suppl 1): 223–31. [523] Stratford TR, Gibbs J, Smith GP. Microstructural analysis of licking behavior following peripheral administration of bombesin or gastrin-releasing peptide. Peptides 1995;16:903–9. [524] Stratford TR, Gibbs J, Smith GP. Simultaneous administration of neuromedin B-10 and gastrin-releasing peptide(1–27) reproduces the satiating and microstructural effects of bombesin. Peptides 1996; 17:107–10. [525] Strohmayer AJ, Smith GP. Cholecystokinin inhibits food intake in genetically obese (C57BL/6j-ob) mice. Peptides 1981;2:39 – 43. [526] Strohmayer AJ, Smith GP. A sex difference in the effect of CCK-8 on food, and water intake in obese (ob/ob), and lean (⫹/⫹) mice. Peptides 1987;8:845– 8. [527] Stuckey JA, Gibbs J, Smith GP. Neural disconnection of gut from brain blocks bombesin-induced satiety. Peptides 1985;6:1249 –52. [528] Suh HW, Kim YH, Choi YS, Song DK. Involvement of different subtypes of cholecystokinin receptors in opioid antinociception in the mouse. Peptides 1995;16:1229 –34.
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Review
Effects of peptides on animal and human behavior: A review of studies published in the first twenty years of the journal Peptides Robert N. McLaya, Weihong Panb,*, Abba J. Kastinb a
b
Naval Medical Center San Diego, Department of Psychiatry, San Diego, CA, USA VA Medical Center and Tulane University School of Medicine, Neuroscience Training Program and Department of Medicine, New Orleans LA 70112-1262, USA Received June 25, 2001; accepted July 27, 2001
Abstract This review catalogs effects of peptides on various aspects of animal and human behavior as published in the journal Peptides in its first twenty years. Topics covered include: activity levels, addiction behavior, ingestive behaviors, learning and memory-based behaviors, nociceptive behaviors, social and sexual behavior, and stereotyped and other behaviors. There are separate tables for these behaviors and a short introduction for each section. © 2001 Elsevier Science Inc. All rights reserved. Keywords: Peptides; Behavior; Activity; Addiction; Ingestion; Learning; Memory; Nociception; Sex; Stereotypy
1. Introduction Why do we do the things we do? This question could be answered on many levels, but increasingly research is discovering ways in which behavior is affected, and sometimes governed, by the actions of peptides. For over twenty years this journal has published articles concerning peptides, many of which involved effects on animal and human behavior. This review is an attempt to catalog those findings. We have focused on studies in which a behavior was elicited after the administration of a peptide. Other types of
studies, such as those studying animals deficient in a particular peptide, or using an agonist to block the effect of a peptide, are for the most part not included, except where the antagonist itself is a peptide or protein. We by no means discount the importance of such studies in understanding how peptides influence behavior, however. Readers wishing to find listings of other effects may be served by reviews previously published in Peptides [3,412,607]. In choosing the articles to include, decisions had to be made concerning what constitutes animal behavior. Usually the choice was obvious. A study of classically conditioned
* Corresponding author. Tel.: ⫹1-504-589-5928; fax: ⫹1-504-522-8559. E-mail address: [email protected] (W. Pan). Abbreviations: 5-HT, Serotonin; 5-HTP, 5-Hydroxytryptophan; 6-OHDA, 6-Hydroxydopamine; A18fa, Ala-Gly Glu-Gly Leu-Ser-Ser-ProPhe-Trp-Ser-Leu-Ala-Ala-Pro-Gln-Arg-Phe-NH2, Admin., Administration; Act., Activity; ACTH, Adrenocorticotropic Hormone; ANF, Atrial Natriuretic Factor; AT, Angiotensin; BBS, Bombesin; BUBUC, Tyr-DCys(StBu)-Gly Phe-Leu-Thr-(OtBu); CCK, Cholecystokinin; CCK-4, Cholecystokinin Tetrapeptide; Tetragastrin; CCK-8, Cholecystokinin Octapeptide; Sincalide; CGRP, Calcitonin Gene-Related Peptide; CRF, Corticotropin Releasing Factor; CTAP, D-Phe-Cys-Try-D-Trp-ORn-Thr-PenThr-NH2; CTOP, D-Phe-Cys-Tyr-D-Tyr-Orn-Thr-NH2; DADELT II, [D-Ala2]Deltorphin; DADLE, D-Ala2-D-Leu5-Enkephalin; DAGO, D-Ala2-MePhe4-Gly-ol5-Enkephalin; DALA, D-Ala2-Methionine-enkephalinamide; DALCE, [D-Ala2,Leu5,Cys6]-Enkephalin; DAMA, D-Ala2Met-Enkephalin Amide; DAME, [D-Ala2-Met5]Enkephalinamide; DAMCK, Tyr-D-Ala-Gly-(NMe)Phe-CH2Cl; DAMGO, Tyr-D-Ala-Gly-
(NMe)Phe-Gly-ol; DDAVP, Desmopressin; 1-deamino-Cys-8-D-Arg-Vasopressin; DGAVP; Desglycinamide-arginine8-Vasopressin; DPEN [D-Pen2,D-Pen5]-Enkephalin; DPDPE, [D-Pen2,D-Pen5]-Enkephalin; DSIP, Delta Sleep-Inducing Peptide; EGF, Epidermal Growth Factor; F8Fa, Phe-Leu-Phe-Gln-Pro-Gln-Arg-Phe-NH2; Frag., Fragment; FMRFa, Phe-Met-Arg-Phe-amide; GHRH, Growth Hormone-Releasing Hormone; GR73632, D-Ala1-[L-Pro9,Me-Leu8]Substance P-(7–11); Ind., Induced; LHRH, Luteinizing Hormone Releasing Hormone; LLTNAM, Lys (CH2NH)-Trp(Nps)-OMe; MCH, Melanin Concentrating Hormone; MIF-1, Melanotropin (MSH) Release-Inhibiting Factor-1; MSH, Melanocyte Stimulating Hormone; NT, Neurotensin; NP-, Neuropeptide-; OT, Oxytocin; Org2766, H-Met-(O2)-Glu-His-Phe-D-Lys-Phe-OH; PACAP, Pituitary Adenylate Cyclase-Activating Polypeptide; PG-KII, pGlu-ProAsn-Pro-Asp-Glu-Phe-Val-Gly-Leu-Met-NH2; PVN, Paraventricular Nucleus; SP, Substance P; TRH, Thyrotropin-Releasing Hormone; TyrMIF-1, Tyr-Pro-Leu-Gly-NH2; Tyr-W-MIF-1, Tyr-Pro-Trp-Gly NH2; VP, Vasopressin; VIP, Vasoactive Intestinal Peptide; YPLG, Tyr-Pro-Leu-Gly
0196-9781/01/$ – see front matter © 2001 Elsevier Science Inc. All rights reserved. PII: S 0 1 9 6 - 9 7 8 1 ( 0 1 ) 0 0 5 5 0 - 2
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eye-blink response is clearly a behavioral study. A study of body temperature is not. In the case of classically conditioned temperature response [65], the choice is less clear. In general, we limited the included studies to those of externally observable actions of an animal or person. Where a study clearly added to the understanding of behavior, however, we were liberal in inclusion. In some cases this may seem somewhat arbitrary. Made only slightly less arbitrary by the existence of authors’ keywords is the decision on how to categorize behavior. We have divided behaviors into seven, broad groups: activity levels, addiction behaviors, ingestive behaviors, learning and memory-based behaviors, nociceptive behaviors, social and sexual behaviors, and stereotyped and other behaviors. We realize that there are many areas of overlap and many ways by which an individual might wish the information organized. Readers with access to the internet and a desire to reorganize the tables to their own liking can find electronic versions of the tables at: http://www.sam.tulane.edu/ classware/pathology/medical_pathology/mcpath/peptides behavior.html For those whose access is limited to the printed page, we hope that these categories are useful. All peptides were administered peripherally unless otherwise noted.
2. Topics 2.1. Activity level Any study of behavior is dependent on the activity level of the organism. Simply put, an animal that is hyperstimulated gives a different response than one that is unconscious or paralyzed. Because of this, motor activity, as measured by distance moved in an enclosed space, is often one of the first effects noted for a peptide. Despite the wealth of data, there is no unifying mechanism yet known to underlie activity level. Some peptides appear to work though common neural substrates such as the ventral striatum and globus pallidus to produce increased activity, whereas others may be independent of such pathways [530]. Still others may regulate activity without directly acting on the brain at all, but rather through peripheral control of metabolism [270]. Peptides appear to be particularly important in the regulation of daily activity and sleep cycles. Balance between neuropeptides such as GHRH and CRF is a key modulator of wakefulness [516]. Other substances such as melatonin are not only believed to be endogenous regulators of sleep, but also have advanced to the clinical stage as pharmaceuticals [192].
Table 1 lists effects on activity published in Peptides. 2.2. Addictive behavior Peptides are critical to the neurobiology of addiction. Often they are the endogenous substrates for the receptors on which drugs of abuse act. The endogenous peptides and their analogs thus have great potential for antagonizing the effects of drugs, and for producing safer, less-addictive versions of addictive substances. Perhaps more importantly, peptide neurotransmitters may underlie certain, central mechanisms of addiction. Drugs of addiction act at a variety of different brain areas and receptors. Common mechanisms of addiction appear to come into play with multiple drugs, however. The mesolimbic dopamine system is the best known [22], but other, peptide-mediated systems also are critical to addiction. Opioid peptide systems within and independent of the mesolimbic system play a role in drug-use reinforcement. CRF and other stress-peptides are critical in withdrawal behaviors, and other peptide neurotransmitters may modulate transmission within the dopamine system [289]. In short, the system is hooked on peptides. Table 2 lists effects on addictive behaviour. 2.3. Nociceptive behavior The mounting of an appropriate response to pain depends on a variety of mechanisms. The painful stimuli must be relayed to the central nervous system. That stimuli must then be appropriately received and a response formed. The signal for the response is then sent back from the nervous system, and the end organ must be capable of carrying out the appropriate action. In some cases pain behaviors can be elicited despite an absence of one or more of these steps. For example, a person might withdraw his hand from an incoming needle because of an expectation of pain even though no pain is or will be experienced. For this review we have relegated such complex behaviors to the learning and memory-related behaviors section. A focus on simple and immediate actions of pain, however, still leaves a number of areas in which peptides influence behavior. The first detector of pain is often referred to as a nociceptor. However, no specific histologic or molecular structure can specifically be said to constitute that receptor. It is believed that noxious stimuli are initially detected by irritation of free, unmyelinated terminal arborizations of nerves. Nociceptive messages are then transmitted by thin myelinated (A-␦) or non-myelinated (C) fibers. A number of substances, including bradykinin, bombesin, galanin, somatostatin, cholecystokinin, vasoactive intestinal peptide, neuropeptide Y, capsaicin, opioids, and cytokines, influence detection and transmission in these nociceptive fibers [43]. Nociceptive signals are carried in the spinal cord via segmental and supraspinal mechanisms. The dorsal horn of
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2183
Table 1 Activity levels Peptide
Movement
Species
Effect
Notes
Vol
Page
Ref
ACTH
Exploratory behavior
Rat
Increased
5
713
546
ACTH ACTH
Motor act. Motor act.
Rat Rat
4 3
117 117
48 48
ACTH frag. (18–24) ACTH frag. (18–24) ACTH frag. (20–24) ACTH/MSH (4–10) analog ebiratide
Sleep Sleep Sleep Sleep
Rat Rat Rat Human
Decreased Increase followed by decrease Increased No effect Increased Decreased
Pre-optic, anterior hypothalamic admin. Chronic admin. Not naltrexone reversible ICV; paradoxical sleep ICV; paradoxical sleep ICV; paradoxical sleep Increased sleep onset latency and increased wakefulness
15 15 15 12
237 237 237 1007
582 582 582 515
ACTH/MSH 4–10 analog ORG 2766 ACTH/MSH 4–10 analog ORG 2766 ACTH/MSH 4–10 analog ORG 2766
Motor act.
Rat
No change
2
255
174
Motor act.
Rat
Increased
First three weeks of life
6
2S 41
1
Motor act.
Rat
Decreased
13
541
235
ACTH/MSH 4–10 analog ORG 2766
Rearing
Rat
Increased
Amygdala admin.; in isolated animals but not controls Rearing and grasping ability occurs earlier in development
6
2S 41
1
ACTH/MSH 4–10 analog ORG 2766 ACTH/MSH frag. (4–10) ACTH/MSH frag. (4–10)
Rearing
Rat
Decreased
5
1061
203
Motor act. Motor act.
Rabbit Rat
No change Increased
6 6
2S 97 285
229 189
ACTH/MSH frag. (4–10) ACTH/MSH frag. (4–10) ACTH/MSH frag. (4–10)
Motor act. Motor act. Motor act.
Rat Rat Rat
Increased Increased Decreased
6 7 9
2S 41 201 277
1 360 593
ACTH/MSH frag. (4–10)
Motor act.
Rat
Increased
9
151
458
ACTH/MSH frag. (4–10)
Rearing
Rat
Increased
6
2S 41
1
Adrenal Peptide E (Frog) Amylin
Motor act. Motor act.
Mouse Rat
No change Decreased
17 17
1291 589
97 91
AT II
Motor act.
Rat
Decreased
9
475
58
BBS BBS BBS
Exploratory behavior Exploratory behavior Motor act.
Rat Rat Rat
Increased Decreased Increased
4 12 4
405 761 693
103 182 355
BBS
Motor act.
Rat
Increased
9
1S 245
260
BBS
Rearing
Rat
Increased
4
693
355
BBS
Sleep
Rat
Decreased
8
237
303
BUBUC
Rearing
Mouse
Increased
11
983
188
In stressed, hypophysectomized rats First three weeks of life In stressed but not controls Neonates; age dependent effect Rearing and grasping ability occurs earlier in development ICV ICV; peptide also antagonized amphetamine-induced hyperactivity; effect blocked by ritanserin ICV; in combination with saralasin but not alone ICV ICV; effect antagonized by neuroleptics With admin. to nucleus accumbens but not nucleus tractus solitarius ICV; effect antagonized by neuroleptics Admin. to multiple brain areas Antagonized by naltrindole
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Table 1 (continued) Peptide
Movement
Species
Effect
Notes
Vol
Page
Ref
Casomorphin ()
Sleep
Rat
Neonates; effect blocked by naloxone
11
1
533
Casomorphin analog BCH 325
Motor act.
Rat
Increased quiet sleep decreased active sleep Decreased
Apomorphine-induced hypoactivity; antagonized by sulpiride
16
635
447
Casomorphin analog BCH 325
Sleep
Rat
12
893
483
Casomorphin derivative BCH 325 CCK
Motor act.
Rat
Decreased at low dose, increased at high dose Increased
13
141
287
Exploratory behavior
Rat
Decreased
4
743
109
CCK
Motor act.
Rat
Decreased
7
587
455
CCK analog Boc[diNle]CCK-7 CCK analog Boc[Phe(2,6diMe)7]-CCK-7 CCK analog Boc-[Phe(2Me)7]-CCK-7 CCK analog Boc-[Phe(4Me)7]-CCK-7 CCK analog Boc[Phe(pentaMe)7]-CCK-7 CCK analog Boc[Phe(triMe)7]-CCK-7 CCK analog Boc-CCK-7 CCK-4 CCK-4
Motor act.
Rat
Decreased
In stress-induced hypoactivity Effect blocked by lesions of nucleus tractus solitarius Effect blocked by vagotomy or capsaicin admin. CNS admin.
6
415
461
Rearing
Mouse
Decreased
ICV
19
301
333
Rearing
Mouse
Decreased
ICV
19
301
333
Rearing
Mouse
Decreased
ICV
19
301
333
Rearing
Mouse
No change
ICV
19
301
333
Rearing
Mouse
Decreased
ICV
19
301
333
Rearing Circling Exploratory behavior
Mouse Rat Rat
Decreased No change Decreased
19 7 16
301 809 815
333 367 215
CCK-4
Motor act.
Rat
Decreased
6
91
275
CCK-4
Rearing
Rat
Decreased
6
91
275
CCK-8 CCK-8
Circling Exploratory behavior
Rat Rat
Increased Decreased
7 5
809 529
367 272
CCK-8
Motor act.
Mouse
Decreased
13
155
231
CCK-8
Motor act.
Rat
No change
4
749
480
CCK-8 CCK-8
Motor act. Motor act.
Rat Rat
Decreased Decreased
6 6
53 91
254 275
CCK-8 CCK-8
Rearing Rearing
Mouse Rat
Decreased Decreased
19 4
301 749
333 480
CCK-8
Rearing
Rat
Decreased
ICV ICV In elevated plus maze, but not in open field. With admin. to nucleus accumbens but not other brain areas With admin. to nucleus accumbens but not other brain areas ICV ICV or IV; antagonized by proglumide ICV or IP; antagonized by CCK-A, but not CCK-B, antagonists Ventral tegmental area and nuclei accumbens septi admin. ICV; in VIP-treated rats With admin. to nucleus accumbens but not other brain areas ICV Ventral tegmental area and nuclei accumbens septi admin. With admin. to nucleus accumbens but not other brain areas
6
91
275
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2185
Table 1 (continued) Peptide
Movement
Species
Effect
Notes
Vol
Page
Ref
CCK-8
Sleep
Mouse
Increased
3
701
610
CCK-8, desulfated Ceruletide analogs Ceruletide analogs
Circling Exploratory behavior Sleep
Rat Mouse Mouse
No change Decreased Increased
Hexobarbital-induced sleep ICV
7 2 3
809 2S 701
367 609 610
CGRP
Motor act.
Rat
Increased
17
1183
294
Corticotropin-like intermediate lobe peptide CRF CRF CRF
Sleep
Rat
Increased
15
237
582
Exploratory behavior Motor act. Motor act.
Monkey Rat Rat
Increased Increased Increased
4 6 6
211 3S 53 923
267 392 161
CRF
Motor act.
Rat
Increased
7
597
476
D-Arg dipeptides
Motor act.
Mouse
Decreased
6
35
475
D-Arg tetrapeptides
Motor act.
Mouse
6
35
475
Dermorphin
Sleep
Chicken
Decrease followed by increase Increased
6
3S 137
397
DSIP
Motor act.
Mouse
Decreased
3
729
211
(1–13) (1–6) (1–7) (1–8)
Motor act. Rearing Circling Circling Circling
Rat Rat and mouse Rat Rat Rat
Biphasic effect No change Increased Increased Increased
3 1 8 8 8
623 341 837 837 837
210 578 186 186 186
Dynorphin frag. (2–17) EGF Endorphin (␣) Endorphin (␣) Endorphin (␣)
Circling Motor act. Motor act. Motor act. Motor act.
Rat Sheep Mouse Mouse Mouse
Increased Decreased No change Decreased Decreased
8 9 13 5 5
837 1221 737 129 1011
186 417 271 555 554
Endorphin (␣) Endorphin ()
Rearing Motor act.
Mouse Goldfish
Decreased Increased
5 3
129 679
555 277
Endorphin () Endorphin () Endorphin ()
Motor act. Motor act. Motor act.
Mouse Rat Rat
No change Decreased Increased
13 1 1
737 179 179
271 603 603
Endorphin () Endorphin () Endorphin ()
Motor act. Motor act. Sleep
Rat Sheep Chicken
Increased Decreased Increased
6 9 6
189 1221 3S 137
601 417 397
Enkephalin (Leu) Enkephalin analog DPDPE Enkephalin analog DPDPE
Circling Motor act.
Rat Rat
Increased Increased
8 13
837 755
186 509
Motor act.
Rat
Increased
14
511
509
DSIP Dynorphin Dynorphin Dynorphin Dynorphin
frag. frag. frag. frag.
Hexobarbital-induced sleep Antagonized by nitric oxide synthase inhibitor ICV; paradoxical sleep
With ICV but not SC ICV; greater effect in controls than in hypophysectomized ICV; blocked by naloxone ICV; mildly antagonized by naloxone ICV; hyperactivity antagonized by naloxone ICV; chicks; effect inhibited by naloxone In amphetamine-treated mice; only at selected doses ICV Substantia nigra admin. Substantia nigra admin. Substantia nigra admin.; not antagonized by naloxone Substantia nigra admin. ICV ICV ICV; not antagonized by naloxone At constant temperatures, but not with temp. gradient
In amphetamine-treated, but not controls ICV ICV; chicks; effect inhibited by naloxone Substantia nigra admin. ICV; morphine-induced hyperactivity ICV; morphine-treated rats
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Table 1 (continued) Peptide
Movement
Species
Effect
Notes
Vol
Page
Ref
analog
Sleep
Chicken
Increased
6
3S 137
397
analog
Motor act.
Rat
Decreased
14
511
509
analog
Motor act.
Rat
Decreased
1
139
257
analog
Motor act.
Human
No change
ICV; chicks; effect inhibited by naloxone ICV; DPDPE and morphine-treated rats ICV; antagonized by taurine but not GABA Elderly
12
871
75
Motor act.
Rat
No change
10
403
443
Mouse
No change
16
1283
556
Galanin
Alternating movements Motor act.
Antagonized morphineinduced horizontal, but not vertical, movement ICV
Rat
No change
11
995
304
Galanin Gastrin I (1–17) GH GHRH GHRH Growth hormone
Motor Motor Motor Motor Motor Motor
act. act. act. act. act. act.
Rat Rat Rat Rat Rat Rat
No change No change Decreased Increased Increased Decreased
Periventricular and PVN admin. PVN admin.
13 1 14 14 14 14
323 383 707 707 708 708
354 408 7 7 7 7
Kassinin-like peptide TK PG-KII L-pyroglutamyl-Lleucine MIF-1
Motor act.
Rat
Increased
18
825
437
Jumping
Rat
Delayed
4
417
295
Circling
Rat
No change
2
189
148
MIF-1
Motor act.
Mouse
1
293
147
MIF-1
Motor act.
Rat
Potentiated increase and suppressed decrease No change
Morphine-withdrawalinduced jumping ICV; dopamine or 5HT stimulated circling Morphine-treated
At 4° or 20°C; not altered by melanin or hypophysectomy
1
243
600
MIF-1 MIF-1
Motor act. Motor act.
Rat Rat
Increased Increased
6 12
189 915
601 293
MIF-1
Motor act.
Mouse
1
293
147
MIF-1 Motilin MSH (␣)
Sleep Running Exploratory behavior
Rat Rat Rat
Potentiated increase and suppressed decrease Increased No change Decreased
6 1 17
7 383 171
440 408 207
MSH (␣)
Motor act.
Rat
No change
1
243
600
MSH (␣) MSH (␣)
Motor act. Motor act.
Rat Rat
No change Increased
2 4
255 865
174 598
MSH (␣) MSH (␣)
Motor act. Motor act.
Rat Rat
Increased Increased
6 16
189 821
601 110
MSH (␣)
Rearing
Rat
Increased
16
821
110
Enkephalin DALA Enkephalin DALCE Enkephalin DAMA Enkephalin DDAVP FMRFa
Galanin
Inhibits habituation Effect with acute but not chronic, treatment ICV
In forced swim test; synergy with tricyclic antidepressants Morphine-induced motor act.
Alcohol-induced sleep Admin. to medial preoptic area; antagonized by MCH At 4° or 20°C; not altered by melanin or hypophysectomy Amphetamine-treated; interactions with L/D cycle and age ICV; antagonized by diazepam or baclofen ICV; antagonized by diazepam or baclofen
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2187
Table 1 (continued) Peptide
Movement
Species
Effect
N-acetylmuramyl-Lalanyl-D-isoglutamine
Sleep (REM)
Rat
Naltrindole
Motor act.
Rat
Decrease followed by increase (no overall, average change) No change
NP-E-I
Motor act.
Rat
Increased
NP-E-I
Rearing
Rat
Increased
NPY NPY NPY NPY
Motor act. Motor act. Resting Sleep cycles
Rat Rat Hamster Hamster
Decreased Decreased Decreased Phase shift
NPY antibody
Circling
Mouse
Increased
NPY antibody
Motor act.
Mouse
Increased
NPY frag. (1–30) NT
Motor act. Motor act.
Rat Guinea pig
Decreased Increased
NT NT NT
Motor act. Motor act. Motor act.
Mouse Mouse Mouse
Increased Decreased Decreased
NT NT NT
Motor act. Motor act. Motor act.
Rat Rat Rat
NT
Motor act.
Rat
No change Decreased Complex interactions with dose and enkephalinase inhibitors Decreased
NT NT NT analog [D-Phe11]-NT NT analog [D-Tyr11]-NT NT analog [Phe11]-NT NT analog Eisai hexapeptide
Rearing Sleep Motor act. Motor act. Motor act. Motor act.
Rat Rat Rat Rat Rat Mouse
No change No change Increased Increased Decreased Decreased
Omega-agatoxin IVA
Motor act.
Rat
No change
Omega-conotoxin GVIA
Motor act.
Rat
No change
Opiate antagonist WIN 44,441–3
Motor recovery
Cat
Increased
Notes
ICV; DPDPE- and morphine-treated rats ICV; antagonized by MSH ICV; antagonized by MSH ICV ICV ICV Geniculohypothalamic tract admin.; animals housed in darkness With admin. to ventromedial thalamus but not other brain areas With admin. to hypothalamus but not other brain areas ICV Acted as noxious stimulus Intrathecal admin. Intrathecal admin. Amphetamine- or phencyclidine-induced hyperactivity PVN and IV admin. ICV
ICV; not antagonized by non-peptide NT antagonist PVN and IV admin. PVN and IV admin.
Amphetamine- or phencyclidine-induced hyperactivity ICV; peptides antagonized druginduced hyperactivity ICV; peptides antagonized druginduced hyperactivity Spinal cord injury
Vol
Page
Ref
5
91
184
14
511
509
18
393
470
18
393
470
15 14 9 15
799 909 1389 1475
54 428 300 245
15
607
579
15
607
579
15 13
799 841
54 454
4 4 18
517 517 1223
249 249 473
4 2 11
493 171 551
511 261 399
16
37
442
4 4 2 2 2 18
493 493 171 171 171 1223
511 511 261 261 261 473
19
1017
404
19
1017
404
6
1S 15
169
(continued on next page)
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Table 1 (continued) Peptide
Movement
Species
Effect
Notes
Vol
Page
Ref
OT
Motor act.
Mouse
Increased
5
535
351
OT Saralasin
Sleep Motor act.
Rat Rat
Increased Decreased
ICV; effect enhanced by co-admin. of amastatin Alcohol-induced sleep ICV; in combination with AT II, but not alone
6 9
7 475
440 58
Sauvagine Sauvagine Sauvagine
Motor act. Motor act. Resting
Mouse Rat Rat
Decreased Increased Increased
6 6 6
3S 59 3S 53 3S 53
420 392 392
Secretin Secretin SK
Motor act. Rearing Motor act.
Rat Rat Rat
Decreased No change Increased
4 4 6
739 739 2S 113
83 83 144
Somatostatin Somatostatin frag. (3–8), (9–14), or (7–10) SP
Rearing Rearing
Rat Rat
Decreased No change
10 10
1153 1153
569 569
Motor act.
Rat
Increased
1
103
262
SP SP
Motor act. Motor act.
Rat Rat
No change Increased
9 6
937 2S 113
173 144
SP frag. (1–7) SP frag. (1–7)
Motor act. Vertical movement
Mouse Rat
Increased Increased
4 20
763 1437
219 126
SP frag. analog pyroglutamylSP(7–11) TRH TRH TRH
Motor act.
Mouse
Decreased
4
763
219
Motor act. Motor act. Motor act.
Rat Rat Rat
Decreased Increased Prevented immobility
2 4 5
131 239 743
602 155 403
TRH
Motor act.
Rat
Increased
6
851
605
TRH
Motor act.
Rat
Increased
5
899
276
TRH analog CG 3704
Motor act.
Rat
Increased
6
851
605
TRH analog DN-1417
Motor act.
Rat
Prevented immobility
5
743
403
TRH analog DN-1417 TRH analog MK-711
Motor act. Motor act.
Rat Rat
Increased Prevented immobility
6 5
1093 743
386 403
TRH analog PS-24 TRH analog RX77386 Urotensin I Urotensin I
Motor act. Motor act. Motor act. Resting
Rat Rat Rat Rat
Increased Increased Increased Increased
2 11 6 6
131 897 3S 53 3S 53
602 258 392 392
Vasotocin Vasotocin
Motor act. Sleep
Cat Cat
Decreased Increased
5 5
25 25
204 204
With ICV but not SC Compared with fasting controls ICV ICV Ventral tegmental area admin. ICV ICV ICV; in controls and haloperidol-treated, but not FLA-63- or methyltyrosine-treated ICV Ventral tegmental area admin. Dorsal periaqueductal gray matter admin.
ICV Depression model; effect present in and after swim, but not before No effect of iron deficiency ICV; effect enhanced by ablation of frontal cortex No effect of iron deficiency Depression model; effect present in and after swim, but not before Depression model; effect present in and after swim, but not before Neonates With ICV but not SC Compared to fasting controls Neonates Neonates
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2189
Table 1 (continued) Peptide
Movement
Species
Effect
Notes
Vol
Page
Ref
Vasotocin
Sleep (quiet state)
Rat
Increased
9
487
230
Vasotocin Vasotocin VIP
Sleep (REM) Sleep (REM) Motor act.
Human Human Rat
Increased Increased Increased
Neonates; dose, sleepstage interactions Narcoleptics Prepubertal boys ICV; enhanced by pargyline, suppressed by CCK-8
1 2 6
281 245 53
418 419 254
VIP VIP
Running Sleep
Rat Rat
Decreased Decreased
1 6
383 53
408 254
VIP VP VP
Sleep (REM) Exploratory behavior Exploratory behavior
Cat Rat Rat
Increased Decreased Increased
5 8 2
837 785 437
154 494 205
VP
Motor act.
Mouse
Increased
5
535
351
VP VP
Motor act. Motor act.
Rabbit Rat
No change Increased
6 15
2S 97 229
229 52
VP
Motor act.
Rat
Decreased
4
717
586
VP
Motor act.
Rat
Decreased
13
17
547
VP
Sleep (slow wave)
Human
Increased
17
1253
424
VP analog DGAVP Z-L-glutamyl-L-leucine
Exploratory behavior Jumping
Rat Mouse
Decreased Delayed
8 4
785 417
494 295
Z-L-glycyl-L-proline
Jumping
Mouse
Delayed
4
417
295
Z-L-leucyl-L-glycine
Jumping
Mouse
No change
4
417
295
Z-L-prolyl-L-leucine
Jumping
Mouse
No change
4
417
295
Z-L-prolyl-D-leucine
Jumping
Mouse
Delayed
4
417
295
ICV; pentobarbitalinduced sleep ICV Also increased latency to move and decreased postural freezing ICV; effect enhanced by co-admin. of amastatin Admin. to neonates, testing in adults In VP-deficient and controls Intrathecal admin.; blocked by V1 antagonist Elderly; subchronic treatment Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping
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Table 2 Addictive behavior Peptide
Behaviour
Species
Effect
ACTH fragment (1–24)
Self-admin. damphetamine Alcohol intake
Rat
Decreased
Rat
Decreased
Aminosenktide
Alcohol-aversive reactions
Rat
Decreased
Bombesin Bombesin
Alcohol intake Alcohol-induced righting impairment Apomorphineinduced yawning Alcohol intake
Rat Rat
Decreased Impaired
Rat
Aminosenktide
Casomorphin analogs CCK-8 CTOP Cyclo(Leu-Gly) Dermorphin analog TDAPA Dermorphin analog TDAPG-NH2 Endorphin () Enkephalin analog DAME Enkephalin (Leu) F8Fa F8Fa F8Fa F8Fa and analogs F8Fa antagonist daY8Ra F8Fa, antibody against FMRa and analogs FMRa and analogs FMRFa FMRFa analog F8Fa Galanin Litorin L-pyroglutamyl-L-leucine
Met-enkephalin-FMRFa chimeric peptide MIF-1
Vol
Page
Ref
5
659
307
19
1557
438
19
1557
438
ICV
6 2
2S 103 1S 99
302 187
Decreased
ICV
13
69
464
Rat
Decreased
14
1193
74
Cocaine-induced seizures Morphine-induced analgesia Morphine tolerance
Rat
Attenuated
Effect blocked by antagonist MK-329 ICV
18
1189
57
Rat
10
767
44
Mouse
Inhibited morphine tolerance No effect
11
139
78
Morphine tolerance
Mouse
No effect
11
139
78
Alcohol-induced righting impairment Ethanol preference and intake Ethanol preference and intake Morphine abstinence syndrome Morphine tolerance
Rat
Impaired
2
1S 99
187
Rat
Decreased
10
697
471
Rat
Decreased
10
697
471
Rat
No change
14
1271
459
Rat
No change
14
1271
459
Morphine withdrawal behavior Morphine abstinence syndrome Morphine abstinence syndrome Morphine abstinence syndrome Morphine abstinence syndrome Morphine abstinence syndrome Morphine withdrawal behavior Morphine withdrawal syndromes Opiate withdrawal behavior Alcohol intake Morphinewithdrawal-induced jumping Morphine tolerance
Rat
Precipitates
11
277
334
Rat
Decreased
12
1011
325
Rat
Decreased
ICV; morphinedependent rats ICV; in F8Fa-treated rats
12
1011
325
Rat
Decreased
11
969
335
Rat
Increased
17
83
484
Rat
Increased
14
731
337
Rat
Precipitates
14
47
338
Rat
Potentiated withdrawal
ICV; morphinedependent rats Third ventricle admin.; morphine-dependent rats Third ventricle admin.; morphine-dependent rats Third ventricle admin.; morphine-dependent rats ICV or IP
20
1211
539
Rat
No change
15
1303
237
Rat Mouse
Decreased Delayed
6 4
2S 103 417
302 295
Mouse
Attenuates tolerance
20
471
214
Rat
Inhibited tolerance
6
7
440
EtOH-induced sleep
Notes
ICV or admin to nucleus basalis magnocellularis; alcohol-preferring rats ICV or admin to nucleus basalis magnocellularis; alcohol-preferring rats; in alcohol-naive but not alcohol-experienced
ICV, intrathecal or SC; antinociceptive effect ICV, intrathecal or SC; antinociceptive effect ICV
Chronic ICV admin. with morphine Chronic ICV admin. with morphine ICV
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Table 2 (continued) Peptide
Behaviour
Species
Effect
MIF-1
Morphine-induced analgesia Morphine abstinence syndrome
Rat Rat
Inhibited morphine tolerance Increased
Alcohol-induced righting impairment Morphine tolerance Alcohol intake Apomorphineinduced yawning Apomorphineinduced yawning EtOH-induced sleep Alcohol intake
Rat
Impaired
Mouse Hamster Rat
Inhibited tolerance Increased Decreased
Rat
Decreased
Rat Rat
Inhibited tolerance Decreased
Alcohol-induced righting impairment Alcohol intake
Rat
Reversed
Rat
Decreased
Rat
Improved
Rat
Decreased
VP
Alcohol-induced righting impairment Apomorphineinduced fighting Alcohol intake
Rat
Decreased
VP (Lys)
Alcohol intake
Rat
Reversed
VP (Lys)
Alcohol pref.
Rat
Reversed
VP analog DGAVP
Alcohol intake
Rat
Reversed
VP analog DGAVP
Alcohol pref.
Rat
Reversed
VP analog DGAVP
Alcohol intake
Rat
Decreased
Z-L-glutamyl-L-leucine
Morphinewithdrawal-induced jumping Morphinewithdrawal-induced jumping Morphinewithdrawal-induced jumping Morphinewithdrawal-induced jumping Morphinewithdrawal-induced jumping
Mouse
Neuropeptide FF
Neurotensin NG-nitro-L-arginine NPY NT NT analog [D-Trp11]NT Oxytocin PG-KII Substance P Tachykinin aminosenkide
TRH TRH
Z-L-glycyl-L-proline
Z-L-leucyl-L-glycine
Z-L-prolyl-leucine
Z-prolyl-D-leucine
Notes
Vol
Page
Ref
10
767
44
17
615
336
2
1S 99
187
ICV
17 9 12
619 1389 755
611 300 398
ICV
12
755
398
6 18
7 825
440 437
1
103
262
19
897
416
2
1S 99
187
9
539
441
4
359
382
6
669
453
6
669
453
6
669
453
6
669
453
6
677
102
Delayed
4
417
295
Mouse
Delayed
4
417
295
Mouse
No change
4
417
295
Mouse
No change
4
417
295
Mouse
Delayed
4
417
295
ICV; antagonized by nitric oxide synthase inhibitors ICV
ICV; alcohol-preferring animals ICV; in haloperidol treated animals Selective effect in lateral hypothalamus and nucleus basalis magnocellularis but not other brain areas ICV
Only in AVP deficient rats AVP deficient with enhanced alcohol intake AVP deficient with enhanced alcohol intake AVP deficient with enhanced alcohol intake AVP deficient with enhanced alcohol intake In roman high avoidance and control rats
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the spinal cord contains many peptides and peptide receptors thought to influence pain, including calcitonin gene related peptide, somatostatin, neuropeptide Y, galanin, and substance P. Opioids also exert one of their main actions at this level by inhibiting release of substance P from primary nociceptive fibers. The spinothalamic and spinoreticular tracts are the best known pathways for transmitting pain from spinal cord to brain, although they are by no means the only course. Several descending pathways also allow the brain to influence the response to pain, and different anatomic locations within the brain influence the response to pain in ways not linked directly to the perception of the stimuli. This presents a plethora of locations at which a peptide can act to influence pain-related behavior within the brain. A listing of pain-related behaviors follows in Table 3. The majority of these studies investigated a simple response to a nociceptive stimuli in the presence of a particular peptide. Therefore rather than listing each paper with the behavior expected described (lifted tail, licked paw, etc.) we have included the protocol used to study the pain behavior and whether the peptide had an analgesic or anti-analgesic effect.
2.4. Ingestive behaviors Regulation of the ingestion of food and water is perhaps the behavior most critical to the survival of an animal. Proper regulation of such behavior depends on communication between brain, gut, and fat tissues, a communication that often makes use of peptides. The OB peptide leptin has received the most attention in recent years, but it should not be forgotten that leptin acts by regulating the expression of other neuropeptides in the brain [2]. These peptides are also active in regulating feeding in pathways that are independent of leptin. Other peripheral satiety systems such as that for CCK [374] are also critical. Drinking, like feeding, is highly regulated by peptides. Osmometric thirst is stimulated by osmoreceptors in the anterior basal forebrain and is influenced by VP. Volumetric thirst on the other hand is regulated by the actions of AT-II in the subfornical organ. Salt appetite also is largely mediated by the AT-II subfornical organ system, although other peptides like CRF and ACTH likewise play a role by stimulating the action of mineralocorticoids on the medial amygdala. Still other peptides appear to simultaneously regulate the intake of both water and food [374], or to regulate intake of particular nutrients [150]. Some nutrients blur the line between food and liquid intake. In particular, alcohol ingestion has often been studied in conjunction with investigations of food and/or water intake. For this reason we present those studies here (Table 4) as well as in the addictive behaviors section.
2.5. Social and sexual behaviors In no field is the idea that biochemicals including peptides, may influence or determine behavior more controversial than in the area of social and sexual behaviors. Although the influence on behavior in humans is often not known, the critical role of peptides in regulating these behaviors in animals is well established. Many of the most well known effects of peptides on sexual and social behaviors involve stimulation of other hormones. Lordosis for example, is said to be controlled by estrogen and progesterone, but estrogen and progesterone secretion in turn is controlled by peptides in the hypothalamus and pituitary gland. More direct effects of peptides on the brain also have been observed. For female sexual behavior the ventromedial nucleus of the hypothalamus plays a particularly important role in peptide receptivity whereas in males the medial preoptic area may be more important [73]. The anatomy of aggressive behavior often centers around the periaqueductal gray matter and the ventral tegmental area, although many studies of the effects of peptides have been done systemically. Social interaction, at least in rodents, may depend on smell, and thus many peptide effects are seen in olfactory areas. Emotional responsiveness provoked in the amygdala is known to be critical to social interactions, and thus the actions of peptides may influence social behaviors in this area. Memory and pain also tie in strongly with social behaviors, which, as mentioned in other sections, leaves quite a wide role for peptides. These behaviours are listed in Table 5. 2.6. Learning and memory behaviors Almost all behaviors are modified by experience. Some behavioral studies, however, are established mainly for the purpose of investigating the effects of experience itself on a particular behavior, in short, how well that behavior is learned. Since peptides act as neurotransmitters and neuromodulators in the brain, it is only to be expected that peptides play a critical role in learning behaviors [487]. The quest for peptides that when exogenously administered alter the ability to perform a learned task has also been the target of much research, since these substances may offer therapeutic hope for dementia [40]. The complexities of behavioral studies of learning can be impressive, teasing out different states of memory, effects on attention, reward, or other motivation. Although the purpose is to investigate internal states, results are gleaned by observation of a particular behavior. For this reason we have organized the learning and memory table by protocol used and, when appropriate, by the particular phase of that protocol in which the behavior was observed (Table 6).
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Table 3 Nociceptive behavior Peptide
Protocol
Species
Effect
Notes
Vol
Page
Ref
ACTH
Ear withdrawal
Rabbit
Anti-analgesic
7
1095
588
AT II
Tail flick
Rat
Analgesic
20
335
445
Calcitonin
Hot plate
Mouse
Analgesic
6
3S 277
200
Calcitonin
Hot plate
Mouse
Analgesic
6
3S 277
200
Calcitonin
Hot plate
Rat
Analgesic
6
3S 273
72
Calcitonin Casomorphin ()
Hot plate Tail flick, hot water, and tail compression test
Rat Rat
Analgesic Analgesic
7 10
315 539
213 585
Casomorphin analogs
Vocalization test
Rat
Analgesic
15
457
465
Casomorphin analogs CCK
Vocalization test Hot plate
Rat Mouse
Analgesic Analgesic
13 18
483 409
466 587
CCK
Tail flick and hot plate
Rat
Anti-analgesic
9
1295
21
CCK analog Boc[Phe(2,6diMe)7]-CCK-7 CCK analog Boc-[Phe(2Me)7]-CCK-7 CCK analog Boc-[Phe(4Me)7]-CCK-7 CCK analog Boc[Phe(pentaMe)7]-CCK-7 CCK analog Boc[Phe(triMe)7]-CCK-7 CCK analog Boc-CCK-7 CCK-8 CCK-8
Hot plate
Mouse
Analgesic
Endorphin ()-induced analgesia ICV; effect antagonized by naloxone ICV, IP, and IV, but no effect SC ICV, IP, and IV, but no effect SC; Ach-induced writhing Intrathecal admin.; antagonized by 6OHDA, haloperidol ICV ICV; greater effect in spontaneously hypertensive rats than controls Effects antagonized by naloxone ICV ICV; analgesic effect antagonized by naloxone but not CCK antagonists Morphine-induced analgesia; effect seen in tail flick but not hot plate ICV
19
301
333
Hot plate
Mouse
Analgesic
ICV
19
301
333
Hot plate
Mouse
Analgesic
ICV
19
301
333
Hot plate
Mouse
No change
ICV
19
301
333
Hot plate
Mouse
Analgesic
ICV
19
301
333
Hot plate Hot plate Hot plate
Mouse Mouse Rat
Analgesic Analgesic Anti-analgesic
19 19 6
301 301 237
333 333 253
CCK-8
Tail flick
Rat
Anti-analgesic
ICV ICV ICV; endorphin-induced analgesia; effect blocked by frontal cortex lesion ICV; morphine- or endorphin-, but not DPDPE-induced, analgesia
16
1229
528
Ceruletide Ceruletide Ceruletide
Mouse Human Human
Analgesic Analgesic Analgesic
2 3 3
2S 61 955 955
132 508 508
Ceruletide analogs Cyclo (Leu-Gly)
Hot plate Pain reporting Tolerance to electricity and thermal pain Hot Plate Tail flick
Mouse Rat
2 10
2S 65 767
609 44
Dalargin
Hot plate
Mouse
Analgesic Inhibited morphine tolerance Analgesic
19
777
482
D-Arg dipeptides
Tail flick
Mouse
Analgesic
6
35
475
Not blocked by naloxone Not blocked by naloxone
Morphine-induced analgesia Carried across the bloodbrain barrier by poly(butyl)cyanoacrylate nanoparticles ICV; mildly antagonized by naloxone
(continued on next page)
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Table 3 (continued) Peptide
Protocol
Species
Effect
Notes
Vol
Page
Ref
Deltakephalin
Tail flick, hot water, and tail compression test
Rat
Analgesic
ICV; greater effect in spontaneously hypertensive rats than controls
10
539
585
Deltorphin Deltorphin II Deltorphin II
Hot plate Tail flick Tail flick
Snail Mice Mouse
Analgesic Analgesic Analgesic
18 17 17
703 1415 619
544 45 611
Deltorphin II analog DADELT II Dermorphin
Tail flick
Rat
Analgesic
13
1123
251
Tail flick
Rat
Analgesic
6
3S 165
59
Tail flick Hot plate and tail flick Tail flick Flinch-jump after foot shock Pain reporting Pain reporting Tail flick Tail flick Hot plate Hot plate Ear withdrawal Hot plate
Rat Rat Mice Rat
Analgesic Analgesic Analgesic Analgesic
ICV; in dermorphin or morphine-tolerant-rats Intrathecal admin. ICV ICV ICV
13 3 17 3
1123 745 1415 41
251 492 45 426
Human Human Rat Rat Slug Slug Rabbit Rat
Analgesic Analgesic Analgesic Anti-analgesic No change No change Analgesic Analgesic
Intrathecal admin. Intrathecal admin. Intrathecal admin. ICV
8 8 6 6 10 10 7 6
191 191 1015 1015 9 9 1095 237
580 580 449 449 116 116 588 253
Hot plate and writhing tests Tail flick Hot plate Hot plate
Mouse
Analgesic
20
865
96
Rat Mouse Slug
Analgesic Analgesic Analgesic
ICV; CCK-8 antagonizes
16 3 10
1229 697 9
528 323 116
Hot plate
Rat
Analgesic
Attenuated by magnetic field
7
449
282
Enkephalin analog DAGO Enkephalin analog DALCE Enkephalin analog DAMA
Hot plate Hot plate Hot plate
Slug Rat Rat
Analgesic Analgesic Analgesic
10 10 1
9 319 139
116 71 257
Enkephalin analog DAMCK
Tail flick
Rat
Analgesic
20
1321
531
Enkephalin analog DPDPE
Hot plate
Snail
Analgesic
19
333
545
Enkephalin analog DPDPE Enkephalin analog DPDPE
Hot plate Tail flick
Snail Rat
Analgesic Analgesic
18 16
703 1229
544 528
Enkephalin analog metkephamide Enkephalin analog DAMGO Enkephalin analog DAMGO FMRFa
Tail flick
Rat
Analgesic
3
869
68
Tail flick
Rat
Analgesic
15
1567
608
Tail flick
Rat
Analgesic
20
1321
531
Hot plate
Rat
No change
10
735
64
Tail flick
Mouse
Analgesic
19
1171
444
Dermorphin Dermorphin analogs DPDPE Dynorphin Dynorphin frag (1–10) Dynorphin frag. (1–13) Dynorphin frag. (1–13) Dynorphin frag. (1–13) Dynorphin frag. (1–17) Dynorphin frag. (1–8) Endorphin () Endorphin () Endorphin () Endorphin () Enkephalin Enkephalin analog DADLE Enkephalin analog DAGO
FMRFa analog [D-Met2]-FMRFa
ICV ICV; not inhibited by NOS inhibitors Intrathecal admin.
ICV ICV; effects enhanced by frontal cortex ablation ICV
ICV ICV; antagonized by taurine and GABA Effect antagonized by naloxone pre-treatment, less so by post-treatment Some cross with pulsedmagnetic field-induced analgesia ICV; CCK-8 does not antagonize Antagonized by naloxone
Effect antagonized by naloxone Morphine-induced analgesia ICV; effect antagonized by antisense to the opioid mu receptor or by antisense to the Gi2␣ G-protein subunit
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Table 3 (continued) Peptide
Protocol
Species
Effect
Notes
Vol
Page
Ref
FMRFa analog A-18Fa
Hot plate
Mouse
Anti-analgesic
12
235
283
FMRFa analog DNS-RFa
Hot plate
Rat
Anti-analgesic
10
735
64
FMRFa analog F8Fa
Hot plate
Mouse
Anti-analgesic
12
235
283
FMRFa analog F8Fa
Hot plate
Mouse
Anti-analgesic
13
603
280
Hemorphin-4
Hot water; tail flick
Mouse
Analgesic
10
747
125
Hemorphin-5
Hot water; tail flick
Mouse
Analgesic
10
747
125
Kentsin
Hot plate and abdominal stretch
Rat
Analgesic
8
613
185
Lys (CH2NH)-Trp(Nps)OMe Lys-Trp
Tail flick (warm water)
Mouse
Analgesic
ICV; defeat-induced analgesia Morphine-induced analgesia ICV; defeat-induced analgesia ICV; morphine- or stress-induced analgesia ICV; naloxone antagonizes ICV; naloxone antagonizes With ICV and intrathecal admin. but not IV; antagonized by naloxone ICV; antagonized by naloxone
13
63
133
Response to heat and electricity Tail flick (warm water)
Mouse and rat
Analgesic
7
39
191
Mouse
Analgesic
13
63
133
20
471
214
Defeat-induced analgesia Defeat-induced analgesia Morphine-treated Rotation-induced analgesia Warm water stressinduced analgesia; no effect with cold waterinduced analgesia Morphine-induced analgesia
6 7 1 10
165 1007 293 493
542 279 147 322
6
1109
190
10
767
44
Morphine-induced analgesia ICV ICV ICis; blocked by GABA mimetics Morphine, but not deltorphin II, tolerance
3
433
121
10 7 8
539 1095 905
585 588 552
17
619
611
7 10 18 4
995 435 1223 517
49 167 473 249
18
1223
473
5 20
747 865
291 96
7 18 9
1095 93 619
588 53 92
3
851
519
Lys-Trp(Nps) Met-enkephalin-FMRFa chimeric peptide MIF-1 MIF-1 MIF-1 MIF-1
Tail flick
Mouse
Analgesic
Hot Hot Hot Hot
Mouse Mouse Mouse Vole
Anti-analgesic Anti-analgesic Anti-analgesic Anti-analgesic
MIF-1
Tail flick
Rat
Anti-analgesic
MIF-1
Tail flick
Rat
MIF-1
Tail flick
Rat
Inhibited morphine tolerance Delayed onset
Morphiceptin MSH Neo-Kyotorphin
Ear withdrawal Tail pinch
Rat Rabbit Mouse
Analgesic No change Analgesic
NG-nitro-L-arginine
Tail flick
Mouse
Inhibited tolerance
N-hexanol-5HTP-5HTP NT NT NT
Tail flick Hot plate Paw pressure test Tail flick
Rat Mouse Mouse Mouse
Analgesic Analgesic Analgesic Analgesic
NT analog Eisai hexapeptide OT Peptide E
Paw pressure test
Mouse
Analgesic
Rat Mouse
Analgesic Analgesic
Somatostatin Somatostatin analog CTAP SP
Tail flick Hot plate and writhing tests Ear withdrawal Tail flick (warm water) Ear withdrawal
Rabbit Mouse Rabbit
No change Analgesic Analgesic
SP
Hot plate
Mouse
Analgesic
plate plate Plate plate
ICV; antagonized by naloxone
ICV ICV Intrathecal admin.; variable effect ICV ICV ICV; only at very high doses ICV ICV; no effect of frag. (1–9) ICV
(continued on next page)
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Table 3 (continued) Peptide
Protocol
Species
Effect
Notes
Vol
Page
Ref
SP
Hot plate
Mouse
Analgesic
Naloxone antagonized effect; SP also antagonized immobilization but not foot shock analgesia
4
763
219
SP SP
Hot plate Motor response and vocalization after electroshock Shock threshold Motor response and vocalization after electroshock Hot plate
Mouse Rat
Analgesic Analgesic
ICV
4 7
31 419
218 390
Cat Rat
Analgesic Analgesic
Intrathecal admin. ICV
5 7
477 419
592 390
Mouse
Analgesic
Naloxone antagonized effect; no effect on stress-induced analgesia ICV
4
763
219
7
419
390
Immobilization but not foot shock analgesia Foot shock-induced analgesia ICV or IV; cold swiminduced analgesia
4
763
219
5
635
69
8
299
79
11
897
258
8
299
70
13
63
133
13
63
133
7 13
1007 1295
279 281
10
493
322
20
971
38
15 15
1508 1567
196 608
5
747
291
6
621
51
7
111
50
SP SP analog DiMe-C7
SP frag. (1–7)
SP frag. (4–11)
Rat
Analgesic
SP frag. (7–11)
Motor response and vocalization after electroshock Hot plate
Mouse
Anti-analgesic
TRH
Tail flick
Rat
Analgesic
TRH
Tail flick
Rat
Analgesic
TRH analog RX 77368
Rat
Analgesic
TRH analog RX 77368
Hot water tail immersion Tail flick
Rat
Analgesic
Trp(Nps)-Lys
Tail flick (warm water)
Mouse
Analgesic
Trp(Nps)(COCH2)(R,S)Lys Tyr-MIF-1 Tyr-MIF-1
Tail flick (warm water)
Mouse
Analgesic
Hot plate Hot plate
Mouse Mouse
Anti-analgesic Anti-analgesic
Tyr-MIF-1
Hot plate
Vole
Anti-analgesic
Tyr-W-MIF-1
Tail flick
Rat
Analgesic
Tyr-W-MIF-1 Tyr-W-MIF-1 and cyclic analogs VP
Tail flick Tail flick
Rat Rat
Analgesic Analgesic
Tail flick
Rat
Analgesic
VP
Tail flick
Rat
Analgesic
VP
Tail flick
Rat
Analgesic
VP VP
Tail flick Tail flick and flinchjump Tail flick
Rat Rat
Analgesic Analgesic
ICV but not SC
13 3
17 613
547 292
Rat
Analgesic
ICV
5
747
291
VP analog DDAVP
ICV; cold swim-induced analgesia ICV; antagonized by naloxone ICV; naloxone antagonizes Defeat-induced analgesia Morphine- or stressinduced analgesia Rotation-induced analgesia ICV; no tolerance, but cross-tolerance with morphine
ICV; not antagonized by naloxone ICV; enhanced by neonatal MSG ICV; paraventricular nucleus lesions block effect
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Table 4 Ingestive behaviors Peptide
Behaviour
Species
Effect
[D-Ser , Leu ]enkephalin-Thr6 Acetorphan
Food intake
Rabbit
Decreased
Milk intake
Cat
Increased
ACTH (1–24)
Food intake
Rat
Decreased
Amylin
Food intake
Mouse
Decreased
Amylin Amylin
Food intake Food intake
Rat Rat
Decreased Decreased
Amylin Amylin
Food intake Food intake
Rat Rat
Decreased Decreased
Amylin
Food intake
Rat
Decreased
Amylin
Food intake
Rat
Decreased
Amylin
Food intake
Rat
Decreased
Amylin
Food intake
Rat
Decreased
Amylin
Saccharin preference
Rat
No change
Amylin Amylin ANF III Anorexigenic peptide Anorexigenic peptide Anorexigenic peptide AT antagonist sarthran AT II AT II
Water intake Water intake Water intake Food intake Food intake Water intake Water intake Salt intake Water intake
Rat Rat Sheep Dog Rat Rat Rat Rat Dog
Decreased No change Decreased Decreased Decreased Decreased Increased Increased Increased
AT AT AT AT AT AT
II II II II II II
Water Water Water Water Water Water
intake intake intake intake intake intake
Hamster Pigeon Rat Rat Rat Rat
Increased Increased Increased Increased Increased Increased
AT AT AT AT
II II II II
Water Water Water Water
intake intake intake intake
Rat Rat Rat Rat
Increased Decreased Increased Increased
Water intake
Rat
Increased
2
AT II
5
Notes
In sham feeding but in not non fasted ICV; also blocked stimulation by opiate agonists ICV or IP; in control, fooddeprived, diabetic, and insulin-treated animals Intra-hypothalamic admin. Intra-hypothalamic admin.; rat amylin more effective than human Vagotomized and controls Effect blocked by histamine antagonists, but not serotonin agonists or antagonists Effect blocked by amylin receptor antagonist In diabetic and control rats; effect antagonized by CGRP antagonist in nondiabetic animals ICV; not influenced by area postrema/nucleus of the solitary tract lesion Effect attenuated by lesion of area postrema/nucleus of solitary tract Intra-hypothalamic admin.; conditioned taste aversion Intra-hypothalamic admin. ICV Sham-feeding ICV infusion in females ICV infusion in females ICV ICV Alkalotic animals unresponsive ICV ICV Attenuated by vagotomy Effect blocked by ANF Greater in spontaneously hypertensive rats Sinoaortic-denervated Subfornical organ lesion Antagonized by ANF Synergistic effects with delta2 opioid agonists ICV; intraoral and free drinking measured
Vol
Page
Ref
7
745
208
10
967
17
7
834
435
12
865
370
13 12
961 919
79 18
16 17
457 1317
326 328
17
119
327
18
643
329
19
1533
330
19
309
331
13
961
79
13 19 12 2 4 4 10 3 9
961 1711 77 2S 235 85 85 261 493 1S 201
79 15 342 288 383 383 223 165 298
4 3 4 6 7 9
103 631 159 1S 145 373 979
358 130 491 212 388 595
10 10 11 18
103 423 837 241
266 265 393 606
19
171
241
(continued on next page)
2198
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 4 (continued) Peptide
Behaviour
Species
Effect
Notes
Vol
Page
Ref
AT II
Water intake
Rat
Increased
19
149
436
AT III
Water intake
Rat
Increased
9
979
595
AT IV
Water intake
Rat
No change
9
979
595
Atrial natriuretic factor Atriopeptin II Avian pancreatic polypeptide BBS BBS
Water intake Water intake Food intake
Rat Rat Chicken
Blocked AT effect Decreased Increased
ICV; effect antagonized by tachykinins Greater in spontaneously hypertensive rats In control and spontaneously hypertensive ICV; AT II-treated AT-stimulated drinking ICV
7 11 9
373 837 449
388 393 141
Alcohol intake Carbohydrate intake
Rat Rat
Decreased Decreased
6 11
2S 103 221
302 346
BBS BBS
Fat intake Feeding interval
Rat Rat
Decreased Decreased
11 12
221 761
346 182
BBS BBS BBS
Food intake Food intake Food intake
Baboon Dog Hamster
Decreased Decreased Decreased
4 5 7
687 675 1079
594 316 24
BBS
Food intake
Mouse
Decreased
9
221
490
BBS BBS BBS BBS BBS
Food Food Food Food Food
intake intake intake intake intake
Rat Rat Rat Rat Rat
Decreased Decreased Decreased Decreased Decreased
2 1 2 2 2
431 131 2S 179 261 409
491 339 198 314 498
BBS BBS
Food intake Food intake
Rat Rat
Decreased Decreased
3 3
61 61
581 581
BBS BBS
Food intake Food intake
Rat Rat
Decreased No change
3 5
553 41
199 589
BBS
Food intake
Rat
Decreased
5
547
39
BBS
Food intake
Rat
Decreased
5
481
94
BBS BBS
Food intake Food intake
Rat Rat
Decreased Decreased
5 6
607 1249
131 527
BBS
Food intake
Rat
Decreased
8
237
303
BBS
Food intake
Rat
Decreased
9
729
128
BBS
Food intake
Rat
Decreased
13
1215
99
BBS BBS
Food intake Food intake
Rat Rat
Decreased Decreased
16 17
903 119
523 327
BBS
Food intake
Rat
Decreased
17
119
327
BBS
Food intake
Rat
Decreased
18
1465
500
BBS
Food intake
Rat
Decreased
19
1407
101
Up to 2 hr post IP admin.; time course different for other macronutrients 4th ventricle admin.; blocked by antagonist In control and vagotomized In short- but not long-day photoperiod Greater effect in old than young
ICV or IP Insulin-induced feeding In controls and vagotomized Ventromedial hypothalamus lesions Admin. to hypothalamus, caudate putamen, olfactory tubercle In control and dorsomedial hypothalamic lesioned Sham-feeding, effect inhibited by proglumide ICV Effect blocked by GI neural disconnect In multiple brain sites; strongest in hypothalamus Nucleus of the tractus solitarius admin. ICV; animals developed tolerance after 8 days Effect partially blocked by amylin receptor antagonist Effect not blocked by amylin receptor antagonist Sham feeding and real feeding Weaker effect in high fat diet
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2199
Table 4 (continued) Peptide
Behaviour
Species
Effect
Notes
Vol
Page
Ref
BBS
Food intake
Rat
Decreased
19
309
331
BBS BBS
Food intake Food intake
Rat Rat
Increased Decreased
20 9
437 1S 233
572 260
BBS BBS BBS
Food intake Food intake Food seeking (bar press) Food seeking (bar press) Food seeking (bar press) Liquid food intake Milk intake
Turkey Wolf Rat
Decreased Decreased Decreased
Effect attenuated by lesion of area postrema/nucleus of solitary tract Amygdala admin. In IP or nucleus tractus solitarius, but not nucleus accumbens, admin. ICV and IV
10 7 12
275 969 1435
140 375 55
Rat
Decreased
13
343
347
Rat
Decreased
4
1
242
Rat Rat
Decreased Decreased
2 12
61 761
301 182
Mouse
Decreased
11
265
181
BBS
Milk seeking (bar press) Protein intake
Rat
Decreased
11
221
346
BBS BBS
Saccharin intake Salt intake
Rat Rat
No change Decreased
2 17
61 951
301 183
BBS BBS BBS BBS BBS BBS
Taste aversion Water intake Water intake Water intake Water intake Water intake
Rat Pigeon Pigeon Rat Rat Rat
No change Increased Increased No change Decreased Decreased
BBS BBS BBS BBS antagonist [D-Phe6, Leu13Cpal4]BBS(6–14) BBS antagonist [D-Phe6, Leu13Cpal4]BBS(6–14) BBS antagonist [D-Phe6, Leu13Cpal4]BBS(6–14) Calcitonin Calcitonin Calcitonin
Water intake Water intake Water intake Food intake
Rat Rat Turkey Rat
Food intake
BBS BBS BBS BBS BBS
Rate-dependent effect Capsaicin-treated and controls
4th ventricle admin.; blocked by antagonist
Up to 30 min post IP admin.; time course different for other macronutrients Sodium-deficient animals; in sham- and normaldrinking conditions
3 3 6 12 2 5
61 631 3S 181 1435 2S 179 41
581 130 129 55 198 589
Decreased Decreased Decreased No change
ICV ICV Rate dependent effect ICV or IP In hypothalamus but not caudate putamen, olfactory tubercle ICV ICV ICV and IV Neonates
5 6 10 14
607 3S 181 275 845
131 129 140 431
Rat
Increased
Only on postnatal day 15
17
1119
356
Water intake
Rat
No change
Neonates
14
845
431
Food intake Food intake Food intake
Dog Hamster Mouse
Decreased Decreased Decreased
In control and vagotomized
5 7 3
675 1079 17
316 24 373
Calcitonin Calcitonin
Food intake Food intake
Mouse Rat
Increased or decreased Decreased
9 7
221 729
490 577
Calcitonin Calcitonin
Food intake Food intake
Rat Rat
Decreased Decreased
3 5
749 861
551 297
Normal and genetically obese, diabetics Age dependent effect In control, lactating, and weaning SC or ICV injection
(continued on next page)
2200
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 4 (continued) Peptide
Behaviour
Species
Effect
Notes
Vol
Page
Ref
Calcitonin
Food intake
Rat
Decreased
9
729
128
Calcitonin
Food intake
Rat
Decreased
Nucleus of the tractus solitarius admin. Effect attenuated by lesion of area postrema/nucleus of solitary tract
19
309
331
Calcitonin Calcitonin Casomorphins Casomorphins
Food intake Water intake Carbohydrate intake Fat intake
Wolf Rat Rat Rat
No change Decreased Decreased Increased
7 3 19 19
969 749 325 325
375 551 319 319
CCK CCK
Carbohydrate intake Fat intake
Rat Rat
Decreased Decreased
11 11
221 221
346 346
CCK CCK CCK
Food intake Food intake Food intake
Dog Hamster Mouse
Decreased Decreased Decreased
5 4 9
675 103 221
316 358 490
CCK
Food intake
Mouse
Decreased
18
1275
341
CCK
Food intake
Rat
Decreased
2
409
498
CCK CCK CCK CCK
Food Food Food Food
intake intake intake intake
Rat Rat Rat Rat
Decreased Decreased Decreased Decreased
3 3 3 5
553 535 535 41
199 108 108 589
CCK
Food intake
Rat
Decreased
5
547
39
CCK
Food intake
Rat
No change
6
1S 133
104
CCK
Food intake
Rat
Decreased
6
927
106
CCK
Food intake
Rat
Decreased
4
15
243
CCK
Food intake
Rat
Decreased
7
587
455
CCK CCK
Food intake Food intake
Rat Rat
Decreased Decreased
8 9
763 601
604 503
CCK
Food intake
Rat
Improved
15
913
158
CCK
Food intake
Rat
Decreased
17
119
327
CCK
Food intake
Rat
Decreased
18
643
329
CCK
Food intake
Rat
Decreased
19
1407
101
CCK
Food intake
Rat
Decreased
19
309
331
CCK
Food intake
Rat
Decreased
1
47
344
ICV but not SC injection ICV or IP; antagonized by enterostatin Up to 6 hr post IP admin.; time course different for other macronutrients In control and vagotomized ICV Greater effect in old than young Synergistic effect with leptin In controls but not vagotomized ICV In hypothalamus but not caudate putamen, olfactory tubercle Effect reduced by dorsomedial hypothalamic lesioned Nucleus tractus solitarius injection Effect blocked by PVN lesions Two-day continuous infusion Effect attenuated by capsaicin Control and VP-deficient Effect blocked by capsaicin in fourth ventricle or vagus, but not spinal cord or pyloric region No effect of sex; effects potentiated by estradiol Effect partially blocked by amylin receptor antagonist In diabetic and control rats; effect antagonized by CGRP antagonist in nondiabetic Weaker effect in high fat diet Effect attenuated by lesion of area postrema/nucleus of solitary tract Hypothalamic or IP; norepinephrine-induced feeding
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2201
Table 4 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
CCK
Rat
Increased
959
264
Rat
Decreased
4
1
242
Mice
No change
2
39
525
CCK
Liquid food intake
Mice
Decreased
2
39
525
CCK
Number of meals
Rat
Decreased
4
15
243
CCK
Protein intake
Rat
Decreased
11
221
346
CCK
Sugar intake
Rat
Decreased
17
593
489
CCK
Time spent eating
Rat
Decreased
4
15
243
CCK
Water intake
Mice
Changed pattern
2
39
525
CCK
Water intake
Rat
Decreased
5
41
589
CCK CCK
Water intake Water intake
Rat Rat
No change No change
8 1
763 47
604 344
CCK agonist devazepide
Food intake
Rat
Increased
In amphetamine-treated but not controls Effect attenuated by food deprivation Normal and geneticallyobese mice Normal and geneticallyobese mice Two-day continuous infusion Up to 1 hr post IP admin.; time course different for other macronutrients Nucleus accumbens admin.; in animals with highbaseline sugar intake, but not in those with lowbaseline intake Two-day continuous infusion Genetically obese mice but not controls In hypothalamus but not caudate putamen or olfactory tubercle Control and VP-deficient Hypothalamic or IP; norepinephrine-induced consumption Effect during estrus, but not diestrus.
16
CCK
Food seeking (bar press) Food seeking (bar press) Inter meal interval
20
451
162
CCK agonist devazepide CCK analog Boc[Phe(2,6diMe)7]CCK-7] CCK analog Boc[Phe(2-Me)7]-CCK-7 CCK analog Boc[Phe(4-Me)7]-CCK-7 CCK analog Boc[Phe(pentaMe)7]CCK-7 CCK analog Boc[Phe(triMe)7]-CCK-7 CCK analog Boc-CCK-7 CCK analog ceruletide CCK analog JMV 170
Meal frequency Food intake
Rat Mouse
No change Decreased
20 19
451 301
162 333
Food intake
Mouse
Decreased
19
301
333
Food intake
Mouse
Decreased
19
301
333
Food intake
Mouse
Decreased
19
301
333
Food intake
Mouse
Decreased
19
301
333
Food intake Food intake Food intake
Mouse Human Rat
Decreased Decreased Decreased
19 3 11
301 607 873
333 506 410
CCK analog JMV 180
Food intake
Rat
Decreased
11
873
410
CCK antagonist L364,718 CCK antagonist L365,260 CCK antagonist L365,260
Food intake
Rat
Increased
40 min IV infusion After ICV but not IP admin. After ICV but not IP admin. CCK-A selective antagonist
12
1215
448
Food intake
Rat
No change
CCK-A selective antagonist
12
1215
448
Sugar intake
Rat
Decreased
IP and nucleus accumbens admin.; CCK-B receptor selective drug; high-sugar consuming rats
17
593
489
CCK
(continued on next page)
2202
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 4 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
CCK antagonist L365,260
Sugar intake
Rat
Increased
17
593
489
CCK frag. CCK-4
Rat
No change
19
849
376
Rat
Induced aversion
ICV, IV, or IP
19
849
376
Rat
Induced conditioning
16
539
Rat
Induced aversion
17
483
377
CCK-8
Taste aversion conditioning Taste aversion conditioning Taste aversion conditioning Taste aversion conditioning Alcohol intake
IP and nucleus accumbens admin.; CCK-B receptor selective drug; low-sugar consuming animals ICV, IV, or IP
Rat
Decreased
14
1193
74
CCK-8 CCK-8
Food intake Food intake
Chicken Dog
Decreased Decreased
3 8
739 651
142 468
CCK-8
Food intake
Hamster
Decreased
7
1079
24
CCK-8 CCK-8
Food intake Food intake
Human Mouse
Decreased
3 8
133 845
507 526
CCK-8 CCK-8 CCK-8 CCK-8
Food Food Food Food
Mouse Rat Rat Rat
Decreased Decreased Decreased Decreased
19 2 2 5
301 431 261 481
333 491 314 94
CCK-8
Food intake
Rat
Decreased
7
729
577
CCK-8
Food intake
Rat
Decreased
7
729
577
CCK-8
Food intake
Rat
Decreased
8
223
495
CCK-8 CCK-8
Food intake Food intake
Rat Rat
Decreased Decreased
9 9
1s 207 1093
481 252
CCK-8 CCK-8
Food intake Food intake
Rat Rat
Decreased Decreased
9 14
989 1193
460 74
CCK-8
Food intake
Rat
Decreased
17
1313
574
CCK-8
Food intake
Rat
Decreased
19
977
89
CCK-8
Food intake
Rat
Decreased
5
97
560
CCK-8 CCK-8
Food intake Food intake
Sheep Sheep
Decreased Decreased
1 3
51 963
137 138
CCK-8 CCK-8
Food intake Food seeking (bar press)
Wolf Rat
Decreased Decreased
7 4
969 351
375 590
CCK-8
Milk seeking (bar press)
Mouse
Decreased
11
265
181
CCK frag. CCK-8 CCK frag. CCK-8 CCK frag. CCK-8
intake intake intake intake
Effect blocked by antagonist MK-329 ICV In ICV but not IV; fooddeprived and insulininduced feeding Greater effect in short-day photoperiod 30 min IV infusion Linear dose relationship in males but not females; greater effect in lean than in obese
Insulin-induced feeding Sham-feeding, effect inhibited by proglumide Greater in lactating mothers than after weaning Resistance to effect during lactation and weaning In control and tumorbearing Reversed by proglumide ICV; proglumide antagonized; no effect from atropine or naloxone NPY treated Effect blocked by antagonist MK-329 In free feeding old and young, but not fixed feeding young. Stronger effect with progesterone and estradiol Tumor-bearing and control animals 6 hr ICV infusion ICV; no effect in cisterna magna Effect altered by GABAergic, dopaminergic and opioid drugs
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2203
Table 4 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
CCK-8 CCK-8
Water intake Water intake
Chicken Dog
Decreased Decreased
3 8
739 651
142 468
CCK-8
Water intake
Mouse
Decreased
8
845
526
CCK-8 CCK-8S Ceruletide
Water intake Food intake Food intake
Mouse Rat Human
Increased Decreased No change
ICV In ICV but not IV; fooddeprived and insulininduced feeding In obese, but not lean, males In lean and obese females
8 17 8
845 355 455
526 576 77
Ceruletide
Food intake
Rat
Decreased
5
1149
499
CGRP
Food intake
Mouse
Decreased
17
511
369
CGRP CGRP
Food intake Food intake
Rat Rat
Decreased Decreased
5 19
861 1533
297 330
CGRP
Mouse
Decreased
17
511
369
CGRP CRF
Milk seeking (bar press) Water intake Food intake
Mouse Rat
No change Decreased
17 4
511 807
369 209
CRF CRF
Food intake Food intake
Rat Rat
Decreased Decreased
6 13
3S 53 879
392 226
CRF
Food intake
Rat
Decreased
20
509
571
CRF
Food seeking (bar press) Food intake
Monkey
Increased
In control and vagotomized, but not adrenalectomized ICV admin. ICV; tail-pinch-induced feeding ICV; not influenced by melanocortin antagonist ICV
10
1199
384
Goldfish
Increased
Serotonin-induced anorexia
19
505
136
Food intake
Rat
ICV; tail-pinch- or NPYinduced feeding
13
879
226
Ethanol preference and intake Food intake Food intake
Rat
Increase at low doses; decrease at high doses Decreased
10
697
471
Goldfish Rat
Increased Increased
17 7
421 711
135 332
Water intake Food seeking (bar press) Water intake Water intake Water intake Food intake Food intake Food intake
Rat Monkey
Increased Increased
18 10
241 1199
606 384
Pigeon Rat Rat Chicken Rat Rat
No change Decreased Increased No change Increased Increased
6 6 18 8 4 7
3S 181 3S 181 241 585 797 711
129 129 606 518 372 332
Food Food Food Food Food
Rabbit Rat Rat Rat Rat and mouse Rat Rat Rat
Increased Increased Increased Increased Increased
7 4 4 4 1
745 797 797 797 341
208 372 372 372 578
4 4 4
797 797 797
372 372 372
CRF antagonist ␣-helical CRF (9–41) CRF antagonist ␣-helical CRF (9–41) DAME DAMGO DADLE Deltorphin II Dermorphin Dermorphin Dermorphin DPDPE Dynorphin Dynorphin Dynorphin Dynorphin Dynorphin Dynorphin Dynorphin Dynorphin
A frag. frag. frag. frag.
(1–10) (1–11) (1–13) (1–13)
Dynorphin frag. (1–8) Dynorphin frag. (1–9) Dynorphin frag. (3–13)
intake intake intake intake intake
Food intake Food intake Food intake
No change No change Increased
Obese women on restricted diet With IP, but not ICV, admin. Not influenced by CCK receptor antagonist ICV; not influenced by area postrema/nucleus of the solitary tract lesion
Admin. to nucleus accumbens septi Interactions with AT II ICV; blocked by naloxone ICV ICV Interactions with AT II ICV; 2 day old ICV Admin. to nucleus accumbens septi
ICV; not reversible by naloxone
(continued on next page)
2204
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 4 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
EGF Eledosin Endorphin () Endorphin () Endorphin ()
Water intake Water intake Food intake Food intake Food intake
Sheep Pigeon Chicken Goldfish Pigeon
Decreased Increased Increased Increased Increased
9 3 9 17 5
1221 631 709 421 691
417 130 345 135 145
Endorphin ()
Food intake
Rat
Increased
3
421
313
Endorphin ()
Food intake
Rat
Increased
7
711
332
Endorphin ()
Water intake
Chicken
9
709
345
Endorphin () Enkephalis analog DAME Enkephalis analog DAME Enkephalis analog DAME Enkephalin (Leu)
Water intake Milk intake Sugar intake Water intake Ethanol preference and intake Sugar intake Water intake Food intake
Pigeon Cat Rat Rat Rat
Decrease at early times, followed by increase at late times No change Decreased No change Decreased Decreased
ICV ICV ICV Mu-receptor effect Pre-fed, ICV injections, only for avian peptide Hypothalamic PVN injection Admin. to nucleus accumbens septi ICV
5 10 10 10 10
691 967 697 697 697
145 17 471 471 471
Rat Rat Rat
No change Decreased Increased
10 10 4
697 697 287
471 471 349
Water intake
Rat
Increased
4
287
349
Food intake
Goldfish
Increased
17
421
135
Carbohydrate intake Fat intake
Rat Rat
No change Decreased
18 18
785 785
409 409
Enterostatin Enterostatin
Fat intake Fat intake
Rat Rat
Decreased Decreased
18 19
657 325
318 319
Enterostatin Enterostatin
Feeding rate Food intake
Rat Rat
Decreased Increased
19 17
557 885
321 451
Enterostatin
Food intake
Rat
18
1341
320
F8Fa
Food intake
Rat
Decreased at low doses but not high doses Decreased
17
353
379
FMRFa
Food intake
Rat
Decreased
6
847
278
Galanin Galanin Galanin
Carbohydrate intake Fat intake Fat preference
Rat Rat Rat
No change No change Increased
18 18 9
207 207 309
496 496 541
Galanin
Food intake
Rat
Decreased
11
995
304
Galanin Galanin Galanin
Food intake Food intake Food intake
Rat Rat Rat
Increased Increased Increased
15 15 17
1303 1267 237
237 497 25
Galanin
Food intake
Rat
Increased
9
309
541
Gastrin
Food intake
Rat
No change
5
41
589
Enkephalin (Leu) Enkephalin (Leu) Enkephalin analog DALA Enkephalin analog DALA Enkephalin analog DPEN Enterostatin Enterostatin
Pre-fed, ICV injection
PVN injection PVN injection; only when food available
ICV ICV; effect blocked by kappa opioid antagonist ICV ICV; antagonized by casomorphins PVN and amygdala admin. ICV; high fat foods; free fed rats Effect when admin. to PVN or amygdala but not other brain areas ICV; no additive effect with naloxone ICV; kappa opiate agonistinduced PVN admin. PVN admin. Paraventricular nucleus admin. After admin. to PVN and periventricular region, but not other hypothalamic areas ICV ICV; effects not sustained Effect blocked by mu antagonists in animals fed high-fat diet Paraventricular nucleus admin. In hypothalamus, caudate putamen, olfactory tubercle
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2205
Table 4 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
Gastrin Gastrin
Water intake Water intake
Rat Rat
Decreased Decreased
Hypothalamic admin. Admin. to caudate putamen, olfactory tubercle
5 5
41 41
589 589
Gastrin (1–17) Gastrin (1–17) Gastrin-related peptide
Food intake Water intake Milk seeking (bar press) Feeding interval Food intake Food intake Food intake Food intake
Rat Rat Mouse
Decreased Decreased Decreased
1 1 11
383 383 265
408 408 181
Rat Rat Rat Rat Rat
Increased Decreased Decreased Decreased Decreased
19 2 3 16 17
1439 2S 179 833 903 107
462 198 517 523 524
Gastrin-releasing peptide
Food intake
Rat
Decreased
18
1465
500
Gastrin-releasing peptide Gastrin-releasing peptide
Food intake Food intake
Rat Rat
No change Decreased
19 20
1439 731
462 16
Gastrin-releasing peptide
Milk intake
Rat
Decreased
20
737
463
Gastrin-releasing peptide
Salt intake
Rat
Decreased
17
951
183
Gastrin-releasing peptide
Sugar intake
Rat
Decreased
20
737
463
Gastrin-releasing peptide GHRH
Water intake Carbohydrate intake
Rat Rat
No change No change
3 15
833 1343
517 150
GHRH
Food intake
Rat
Increased
15
1343
150
GHRH
Food intake
Rat
Increased
16
595
558
GHRH
Food intake
Rat
Decreased
2
261
314
GHRH
Food intake
Rat
9
1S 35
557
GHRH
Protein intake
Rat
Increase at low doses; decrease at high doses Increased
15
1343
150
GHRH, antibody to
Carbohydrate intake
Rat
No change
16
7
149
GHRH, antibody to
Protein intake
Rat
Decreased
16
7
149
Glucagon
Food intake
Dog
Decreased
5
675
316
Glucagon Glucagon
Food intake Food intake
Hamster Mouse
No change Decreased
7 9
1079 221
24 490
Glucagon Glucagon Glucagon
Food intake Food intake Food intake
Rat Rat Rat
Decreased Decreased Increased
3 3 8
163 163 943
195 195 194
Gastrin-releasing Gastrin-releasing Gastrin-releasing Gastrin-releasing Gastrin-releasing
peptide peptide peptide peptide peptide
Synergistic effect with neuromedin B In real feeding but not sham feeding Sham feeding; Synergistic effect with pre-feeding Effect on free feeding but not intraoral intake Sodium-deficient animals; in sham- and normaldrinking Effect on free feeding but not intraoral intake Suprachiasmatic nucleus/ medial preoptic area admin. Suprachiasmatic nucleus/ medial preoptic area admin.; effect blocked by methylnaltrexone Suprachiasmatic nucleus admin.; light-cycle dependent effect Hypothalamic admin.; insulin-induced feeding ICV Suprachiasmatic nucleus/ medial preoptic areas admin.; effect blocked by methyl-naltrexone Suprachiasmatic/preoptic area admin.; feeding studied at dark onset Suprachiasmatic/preoptic area admin.; feeding studied at dark onset In control, but not vagotomized Greater effect in old than young Sham-feeding
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2206
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Table 4 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
Glucagon
Food intake
Rat
Decreased
Effect blocked by amylin receptor antagonist when given at dark onset
17
119
327
Glucagon frag. (1–21) Glucagon-like peptide-1(7–37) amide Insulin Insulin Insulin
Food intake Food intake
Rat Rat
No change Decreased
ICV; in lean and obese
8 19
943 869
194 248
Food intake Food intake Food intake
Rat Rat Rat
Streptozocin-diabetic rats
4 12 13
79 425 721
124 98 348
Insulin Interleukin-1
Food intake Food intake
Rat Rat
Increased Increased Decreased (returned to same as controls) Decreased Decreased
19 14
1549 175
156 553
Interleukin-1
Food intake
Rat
Decreased
17
517
502
Kassinin
Water intake
Rat
Decreased
10
125
422
Leptin Leptin
Food intake Food intake
Mouse Mouse
Decreased Decreased
20 18
595 1275
368 341
Leptin Litorin Litorin Litorin Litorin Litorin Litorin MIF-1
Food intake Alcohol intake Food intake Food intake Water intake Water intake Water intake Food intake
Rat Rat Rat Rat Pigeon Rat Rat Rat
Decreased Decreased Decreased Decreased Increased Decreased Decreased No change
19 6 2 5 6 6 5 15
1549 2S 103 2S 179 607 3S 181 3S 181 607 243
156 302 198 131 129 129 131 216
MIF-1 Motilin
Water intake Food intake
Rat Mouse
Decreased Increased
1 19
353 987
407 13
Motilin Motilin Motilin MSH (␣)
Food intake Food intake Water intake Food intake
Rat Rat Rat Rat
Decreased Increased Decreased Decreased
1 6 1 7
383 41 383 834
408 193 408 435
MSH frag. analog Lys-D-Pro-Thr MSH frag. Lys-Pro MSH frag. Lys-Pro-Val
Food intake
Rat
Decreased
14
175
553
Food intake Food intake
Rat Rat
Increased Increased
14 14
175 175
553 553
Neoendorphin (␣)
Food intake
Rat
Increased
7
711
332
Neurokinin A
Water intake
Rat
Decreased
11
339
423
Neuromedin B
Food intake
Rat
Decreased
17
107
524
Neuromedin B
Salt intake
Rat
No change
17
951
183
NPY NPY
Alcohol intake Amino acid deficient diet intake
Hamster Rat
Increased Decreased
9 19
1389 527
300 111
Hypoglycemic rats Antagonized by MSH frag. Lys-Pro-Val and frag. analog Lys-D-Pro-Thr ICV; effect blocked by NPY Neonates; AT II-induced thirst Genetically obese mice Synergistic effect with CCK Hypoglycemic rats
ICV ICV ICV ICV Peptide YY-induced hyperphagia ICV; attenuated by motilin receptor antagonist Fasted subjects ICV; also blocked stimulation by opiate agonists In IL-1-induced anorexia but not controls In IL-1-induced anorexia In IL-1-induced anorexia but not controls Admin. to nucleus accumbens septi Neonates; in control and AT-treated Synergistic effect with gastrin releasing peptide Sodium-deficient animals; in sham- and normaldrinking Anterior piriform complex admin.; threonineimbalanced model
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R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2207
Table 4 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
NPY NPY NPY
Carbohydrate intake Carbohydrate intake Carbohydrate intake
Rat Rat Rat
Increased Increased Increased
6 12 18
1205 1251 207
510 312 496
NPY NPY NPY
Fat intake Fat intake Food intake
Rat Rat Chicken
No change No change Increased
12 18 8
1251 207 585
312 496 518
NPY NPY
Food intake Food intake
Chicken Dog
Increased Decreased
Admin. to PVN PVN admin. PVN admin.; in carbohydrate-preferring, but not control, rats PVN admin. PVN admin. ICV; 2 day old; in animals that did not have convulsions ICV ICV; in insulin-treated, but not fasted or satiated
8 8
823 651
299 468
NPY NPY NPY
Food intake Food intake Food intake
Hamster Mouse Rat
Increased Increased Increased
9 10 5
1389 963 1025
300 177 315
NPY NPY
Food intake Food intake
Rat Rat
Increased Increased
7 9
1189 723
512 269
NPY
Food intake
Rat
Increased
9
1269
467
NPY
Food intake
Rat
Increased
9
989
460
NPY
Food intake
Rat
Increased
10
1283
81
NPY
Food intake
Rat
Increased
12
609
411
NPY NPY NPY
Food intake Food intake Food intake
Rat Rat Rat
Increased Increased Increased
12 13 14
1251 581 475
312 513 514
NPY NPY
Food intake Food intake
Rat Rat
Increased Increased
15 16
799 599
54 112
NPY
Food intake
Rat
Decreased
17
797
80
NPY
Food intake
Rat
Increased
17
797
80
NPY
Food intake
Rat
Increased
17
797
80
NPY
Food intake
Rat
Increased
17
517
502
NPY
Latency to onset of feeding Milk seeking (bar press) Protein and fat intake Protein intake Sugar intake Sugar intake Water intake
Rat
Decreased
9
1269
467
Mouse
Increased
12
1329
178
Rat Rat Rat Rat Chicken
No change Decreased Decreased Increased No change
6 12 20 20 8
1205 1251 601 601 823
510 312 543 543 299
NPY NPY NPY NPY NPY NPY
ICV Effect inhibited by naloxone and haloperidol PVN of hypothalamus ICV; satiated females; attenuated by adrenalectomy; no effect from CCK ICV; effect attenuated by hypothalamic transection ICV; attenuated by intravenous infusions of glucose, but not fructose Paraventricular hypothalamic admin.; effect blocked by pertussis toxin Sustained admin. to PVN of hypothalamus PVN admin. Hypothalamic admin. Hypothalamic perifornical admin.; similar effects at different times of day ICV Medial PVN and lateral perifornical hypothalamus admin. Hypothalamic admin.; tumor-bearing Hypothalamic admin.; constant infusion; tumorbearing Daily increasing doses; effect in control but not tumor-bearing ICV; in IL-1-treated and controls ICV; effect attenuated by hypothalamic transection ICV; also overcame taste aversion Admin. to PVN PVN admin. ICV; continuous infusion ICV; single admin. ICV
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2208
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Table 4 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
NPY
Water intake
Dog
Decreased
ICV; in insulin-treated, but not fasted or satiated
8
651
468
NPY NPY
Water intake Water intake
Hamster Rat
Increased Increased
9 5
1389 1025
300 315
NPY analog [Leu31, Pro34]NPY NPY analog [Leu31, Pro34]NPY NPY analog [Leu31, Pro34]NPY NPY analog [Leu31, Pro34]NPY NPY analog C2-NPY
Carbohydrate intake
Rat
Increased
12
1251
312
Fat intake
Rat
No change
12
1251
312
Food intake
Rat
Increased
12
1251
312
Protein intake
Rat
Decreased
12
1251
312
Food intake
Rat
13
581
513
NPY analog [Pro34]NPY NPY antagonist BW1229U91 NPY antibodies NPY antibody
Food intake Food intake
Rat Rat
Increased (slight effect) Increased No change
Analog is Y1 selective ICV; control and obese
13 20
581 367
513 583
Food intake Food intake
Rat Rat
Decreased Decreased
13 15
581 607
513 579
NPY frag. (1–30) NPY frag. (13–36)
Food intake Carbohydrate intake
Rat Rat
No change Decreased
15 12
799 1251
54 312
NPY frag. (13–36)
Food intake
Rat
Decreased
12
1251
312
NPY frag. (13–36)
Protein intake
Rat
Decreased
12
1251
312
NPY NPY NPY NPY NPY NPY NPY NPY NT
Food intake Carbohydrate intake Fat intake Food intake Food intake Protein intake Food intake Food intake Food intake
Mouse Rat Rat Rat Rat Rat Mouse Rat Rat
No change Increased No change Increased Increased Increased No change No change Decreased
10 12 12 12 13 12 10 13 4
963 1251 1251 1251 581 1251 963 581 493
177 312 312 312 513 312 177 513 511
NT
Food intake
Rat
Decreased
9
729
128
NT
Water intake
Rat
No change
4
493
511
OT
Food intake
Rat
Decreased
10
89
12
OT
Food intake
Rat
Decreased
12
113
405
OT OT
Food intake Food intake
Rat Rat
Decreased Decreased
17 17
1119 1119
356 356
OT
Latency to first meal
Rat
Increased
ICV In ventromedial hypothalamus and ventromedial and ventrolateral thalamus, but not other hypothalamic areas ICV PVN admin.; frag. is Y2 selective PVN admin.; frag. is Y2 selective PVN admin.; frag. is Y2 selective ICV PVN admin. PVN admin. PVN admin. Hypothalamic admin. PVN admin. ICV Hypothalamic admin. Hypothalamic PVN injection Nucleus of the tractus solitarius admin. Hypothalamic PVN injection IP or ICV; blocked by pretreatment with d(CH2)5Tyr(Me)-Orn8vasotocin Antagonist blocked effect, but had no effect on its own In neonates and adults In neonates and adults; effect greater in adults IP or ICV; blocked by pretreatment with d(CH2)5Tyr(Me)-Orn8vasotocin
10
89
12
frag. (20–36) frag. (2–36) frag. (2–36) frag. (2–36) frag. (2–36) frag. (2–36) frag. (26–36) free acid
Effect inhibited by naloxone and haloperidol PVN admin.; analog is Y1 selective PVN admin.; analog is Y1 selective PVN admin.; analog is Y1 selective PVN admin.; analog is Y1 selective Analog is Y2 selective
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2209
Table 4 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
OT
Time spent eating
Rat
Decreased
10
89
12
OT antagonist vasotocin PACAP PACAP
Food intake Food intake Food intake
Rat Mouse Rat
Increased Increased Decreased
17 13 16
1119 1133 1511
356 371 82
PACAP
Water intake
Rat
Decreased
16
1511
82
Pancreatic polypeptide Pancreatic polypeptide Pancreatic polypeptide
Food intake Food intake Food intake
Chicken Dog Human
Increased No change Decreased
IP or ICV; blocked by pretreatment with d(CH2)5Tyr(Me)-Orn8vasotocin Only on postnatal day 10 ICV Hypothalamic admin.; in NPY-treated and controls Hypothalamic admin.; in NPY-treated and controls Intracerebral inj.; 2 day old
8 5 14
585 675 497
518 316 42
Pancreatic polypeptide Pancreatic polypeptide Peptide YY Peptide YY Peptide YY
Food intake Food intake Carbohydrate intake Food intake Food intake
Rat Rat Rat Chicken Rat
Increased Decreased Increased Increased Increased
11 20 6 8 15
673 1445 1205 823 243
19 14 510 299 216
Peptide YY Peptide YY PG-KII
Protein and fat intake Water intake Alcohol intake
Rat Chicken Rat
No change No change Decreased
6 8 18
1205 823 825
510 299 437
PG-KII
Food intake
Rat
No change
18
825
437
PG-KII
Water intake
Rat
Decreased
18
825
437
Ranatensin Ranatensin Ranatensin Ranatesin Sauvagine
Water intake Water intake Water intake Food intake Food intake
Pigeon Rat Rat Rat Rat
Increased Decreased Decreased Decreased Decreased
6 6 5 5 6
3S 181 3S 181 607 607 3S 53
129 129 131 131 392
Somatostatin
Amino acid deficient diet intake
Rat
Biphasic time and dose-related-effect
19
527
111
Somatostatin Somatostatin Somatostatin Somatostatin analog SMS 201–995 Somatostatin antibody Tachykinin agonist NP-␥
Food Food Food Food
Dog Rat Rat Rat
No change Decreased Increased Increased
5 19 19 9
675 991 991 1S 211
316 477 477 117
Food intake Water intake
Rat Rat
Decreased Decreased
9 19
1S 211 149
117 436
Tachykinin aminosenkide
Alcohol intake
Rat
Decreased
19
897
416
Tachykinin aminosenkide
Food intake
Rat
Decreased
19
897
416
Tachykinin aminosenktide
Water intake
Rat
Decreased
19
897
416
intake intake intake intake
Prader-Willi syndrome; effect only seen in females Hypothalamic admin. ICV Admin. to PVN ICV ICV; antagonized by naloxone Admin. to PVN ICV ICV; alcohol-preferring animals ICV; alcohol-preferring animals ICV; alcohol-preferring animals; in AT-stimulated drinking, but not control, animals ICV ICV ICV ICV ICV, and SC in vagotomized animals Anterior piriform complex admin.; Threonineimbalanced model In dark phase In light phase ICV ICV ICV; AT-stimulated drinking Selective effect in lateral hypothalamus and nucleus basalis magnocellularis but not other brain areas Effect with admin. to stria terminalis but not other brain areas Effect with admin. to stria terminalis but not other brain areas
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Table 4 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
Tachykinin analog Suc-[Asp6,MePhe8]Substance P(6–11) Tachykinin analog Suc-[Asp6,MePhe8]Substance P(6–11) Tachykinins Tachykinins TRH Tuftsin
Food intake
Rat
Decreased
Alcohol-preferring animals
16
533
88
Food intake
Rat
Alcohol-preferring animals
16
533
88
Water intake Water intake Food intake Food seeking (bar press) Food intake
Pigeon Rat Dog Monkey
Decreased at low doses but not high doses Increased Decreased Decreased Increased
ICV ICV Sham-feeding ICV
6 6 2 10
3S 181 3S 181 2S 235 1199
129 129 288 384
Rabbit
Decreased
7
745
208
Food intake
Rat
Increased
20
1425
114
Tyr-MIF-1
Food intake
Rat
No change
15
243
216
Urotensin VIP VIP VP VP
Food intake Food intake Water intake Alcohol intake Water intake
Rat Rat Rat Rat Rat
6 1 1 4 12
3S 53 383 383 359 285
392 408 408 382 584
VP
Water intake
Rat
Decreased No change No change Decreased Decreased (returned to same as controls) Decreased
1
11
190
VP VP (Lys)
Water intake Alcohol intake
Sheep Rat
Decreased Reversed
9 6
1221 669
417 453
VP (Lys)
Alcohol preference
Rat
Reversed
6
669
453
VP analog DGAVP
Alcohol intake
Rat
Decreased
6
677
102
VP antagonists VP frag. DGAVP
Water intake Alcohol intake
Dog Rat
Decreased Reversed
8 6
1003 669
532 453
VP frag. DGAVP
Alcohol preference
Rat
Reversed
6
669
453
Tyr-D-Ala-Gly-(Me) Phe-Gly-ol Tyr-MIF-1
Neonates; stress-induced feeding Peptide YY-induced hyperphagia ICV admin.
Only in VP-deficient rats Food restricted; VPdeficient Hypothalamic graft of VPproducing cells in VPdeficient rats ICV VP-deficient with enhanced alcohol intake VP-deficient with enhanced alcohol intake In Roman high avoidance and control rats ICV VP-deficient with enhanced alcohol intake VP-deficient with enhanced alcohol intake
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2211
Table 5 Social and sexual behavior Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
A18Fa octa-decapeptide
Defeat behavior
Mouse
Increased
12
235
283
A18Fa, antibody to
Defeat behavior
Mouse
Decreased
12
235
283
ACTH
Sexual behavior
Newt
4
729
361
ACTH ACTH frag. (4–10) ACTH/MSH 4–10 analog ORG 2766 ACTH/MSH 4–10 analog ORG 2766 Amylin
Sexual behavior Sexual behavior Social interaction
Rat Newt Rat
Increased ICV; decreased IP Decreased Increased Increased
ICV; subordinate mice; possible anti-analgesic effect ICV; subordinate mice; possible analgesic effect Opposite effects ICV or IP
14 4 2
379 729 255
8 361 174
Social interaction
Rat
Increased
2
255
174
Sexual behavior
Rat
Decreased
20
379
90
CCK-4
Defensive behavior
Rat
Increased
20
383
575
CGRP
Sexual behavior
Rat
Increased
20
379
90
CRF CRF CRF Dynorphin Endorphin ()
Aggressive behavior Defensive behavior Social interaction Lordosis Lordosis
Mouse Mouse Mouse Rat Rat
8 8 8 8 8
935 935 935 309 309
353 353 353 427 427
Enkephalin (Met)
Aggressive behavior
Goldfish
Decreased Increased Decreased No change Dose-dependent increase or decrease Decreased
1
73
474
Enkephalin (Met)
Aggressive behavior
Fish
6
139
86
Enkephalin analog [D-Ala2-Met5]enkephalinamide
Defensive behavior
Cat
9
999
486
F8Fa
Defeat behavior
Mouse
Increased
12
235
283
F8Fa, antibody to
Defeat behavior
Mouse
Decreased
12
235
283
LHRH
Sexual behavior
Human
No change
2
1S 115
163
LHRH antagonist MCH
Sexual behavior Sexual behavior
Rat Rat
Decreased Increased
2 17
393 171
157 207
MIF-1 MIF-1 Morphiceptin
Aggressive behavior Aggressive behavior Lordosis
Mouse Mouse Rat
ICV
6 7 8
165 1007 309
542 279 427
MSH
Lordosis
Rat
ICV or SC
7
581
446
MSH (␣)
Aggressive behavior
Mouse
Decreased Decreased Dose-dependent inhibition or facilitation Increased or decreased Increased
1
69
400
MSH (␣)
Aggressive behavior
Rat
Increased
17
171
207
MSH (␣)
Sexual behavior
Newt
Increased
Aggression seen in mouse paired with MSHtreated animal Admin. to ventromedial nucleus; antagonized by MCH ICV
4
729
361
Increased at low dose, decreased at high dose Increased at low dose, decreased at high dose
ICV In familiar, but not in unfamiliar, surroundings
ICV; effect blocked by CGRP Ultrasound-induced defense behavior ICV; in rats with amylinsuppressed sexual behavior ICV ICV ICV ICV ICV ICV; interaction with melatonin
Admin. to midbrain periaqueductal gray; electrical stimulationelicited defense behavior ICV; subordinate mice; possible anti-analgesic effect ICV; subordinate mice; possible analgesic effect Married men with secondary impotence ICV Admin. to ventromedial nucleus or medial preoptic area
(continued on next page)
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Table 5 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
MSH (␣)
Sexual behavior
Rat
Increased
Admin. to ventromedial nucleus or medial preoptic area
17
171
207
MSH (␣) MSH (␣) MSH/ACTH 4–10 analog Org2766 OT OT
Social interaction Social interaction Social interaction
Mouse Rat Rat
No change Decreased No change
2 2 13
1S 123 255 541
105 174 235
Human Rat
No change Preserved recognition
13 19
461 999
62 151
OT Sauvagine Sauvagine Sauvagine Somatostatin SP SP
Facial recognition test Social discrimination response Social recognition Aggressive behavior Defensive behavior Social interaction Aggressive behavior Aggressive behavior Aggressive behavior
Rat Mouse Mouse Mouse Fish Fish Mouse
Increased Decreased Increased Decreased Decreased Decreased Decreased
19 8 8 8 6 6 5
999 935 935 935 139 139 86
151 353 353 353 86 86 220
SP frag. (1–7) SP frag. (1–7)
Aggressive behavior Aggressive behavior
Mouse Mouse
Decreased Decreased
4 5
763 86
219 220
SP frag. (4–11)
Aggressive behavior
Mouse
No change
5
86
220
SP frag. analog pyroglutamylSP(7–11) SP frag. analog pyroglutamylSP(7–11) TRH
Aggressive behavior
Mouse
Increased
4
763
219
Aggressive behavior
Mouse
Increased
5
86
220
Dominance
Rat
Increased
9
539
441
TRH
Fighting
Rat
Decreased
9
539
441
TRH TRH analog RGH 2202 Tyr-MIF-1 Vasotocin VP
Sexual behavior RGH 2202 Sexual behavior Aggressive behavior Aggressive behavior Lordosis
Rat Rat
Increased Increased
12 12
1309 1309
152 152
Mouse Rat Hamster
Decreased Decreased Increased
7 5 14
1007 1135 1049
279 61 244
VP VP
Sexual behavior Social discrimination response Social interaction Social recognition Facial recognition test
Newt Rat
Increased Preserved recognition
4 19
97 999
365 151
Mouse Rat Human
No change Increased No change
2 19 13
1S 123 999 461
105 151 62
VP VP VP analog DGAVP
Admin. to central nucleus of amygdala Olfactory bulb admin. Olfactory bulb admin. ICV ICV ICV
Isolation-enhanced aggression Isolation-enhanced aggression; effect enhanced by naloxone Isolation-enhanced aggression
Isolation-enhanced aggression; effect antagonized by naloxone Admin. to nucleus accumbens septi; in dominant, but not subordinate, animals Admin. to nucleus accumbens septi; shockinduced and apomorphine-induced fighting
Pinealectomized animals Medial preoptic area admin. ICV or IP
Olfactory bulb admin.
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2213
Table 6 Learning and memory behaviors Peptide
Paradigm
Phase
Species
ACTH ACTH ACTH
Active avoidance Acquisition and recall Mouse Active avoidance Acquisition and recall Mouse Passive avoidance response Recall Rat
ACTH
Thomson Box visual discrimination Thomson Box visual discrimination Thomson Box visual discrimination Active avoidance
Effect
Notes
Vol Page
Ref
Acquisition
Rat
Improvement Given prenatally 9 Improvement Given postnatally 9 Decreased avoidance 10 latency No improvement 1
Extinction
Rat
No improvement
1
277
472
Reversal
Rat
No improvement
1
277
472
Extinction
Rat
Delayed
ICV
2
389
570
Passive avoidance response Acquisition
Rat
Improvement
ICV
2
389
570
Thomson Box visual discrimination Thomson Box visual discrimination Thomson Box visual discrimination Active avoidance
Acquisition
Rat
No improvement
1
277
472
Extinction
Rat
No improvement
1
277
472
Reversal
Rat
No improvement
1
277
472
Retention
Rat
Improvement
9
575
223
ACTH analog Ebiratide
Passive avoidance behavior Retention
Rat
Improvement
9
575
233
ACTH analog Ebiratide
Passive avoidance behavior Retention
Rat
Improvement
9
575
233
ACTH analog Ebiratide
Radial Maze (8 arm)
Working memory
Rat
Improvement
9
575
233
ACTH analog Ebiratide ACTH analog HOE 427
Retention Acquisition
Rat Rat
Improvement No change
9 10
575 303
233 456
Recall
Rat
No change
10
303
456
Reversal
Rat
Impaired reversal
10
303
456
ACTH analog ORG 2766 ACTH analog ORG 2766
Up-hill avoidance task Two-smell discrimination task Two-smell discrimination task Two-smell discrimination task Active avoidance Active avoidance
Acquisition and recall Mouse Acquisition and recall Mouse
Improvement Improvement
9 9
745 745
239 239
ACTH analog ORG 2766
Active behavior response
Extinction
Rat
Slowed
7
563
172
ACTH analog ORG 2766
Morris water maze
Recall
Rat
Improvement
11
911
432
ACTH analog ORG 2766
Passive avoidance response Recall
Rat
No change
11
911
432
ACTH analog ORG 2766 ACTH analog ORG 2766
Passive behavior response Passive behavior response
Acquisition Acquisition
Rat Rat
No change Improved
7 7
563 563
172 172
ACTH analog ORG 2766 ACTH analog ORG 2766
Passive behavior response Passive behavior response
Retention Retention
Rat Rat
Improved Improved
7 7
563 563
172 172
ACTH analog ORG 2766 ACTH analog ORG 2766 ACTH analog ORG 2766
Acquisition Extinction Acquisition
Rat Rat Rat
Improved No change No change
6 6 10
2S 41 2S 41 303
1 1 456
Recall
Rat
Improvement
10
303
456
ACTH/MSH frag. (4–10)
T-maze T-maze Two-smell discrimination task Two-smell discrimination task Active avoidance
Acquisition and recall Mouse
Improvement
745
239
ACTH/MSH frag. (4–10)
Active avoidance
Acquisition and recall Mouse
No change
745
239
ACTH ACTH ACTH analog -(Tyr9) melanotropin (9–18) ACTH analog -(Tyr9) melanotropin (9–18) ACTH analog D-PheMSH-ACTH ACTH analog D-PheMSH-ACTH ACTH analog D-PheMSH-ACTH ACTH analog Ebiratide
ACTH analog HOE 427 ACTH analog HOE 427
ACTH analog ORG 2766
Socially-deprived animals ECT-induced amnesia Scopolamineinduced amnesia Scopolamineinduced amnesia
Given prenatally Given postnatally; in males only Phase-shiftinduced amnesia Rats with fimbria fornix transection Rats with fimbria fornix transection Phase-shiftinduced amnesia Phase-shiftinduced amnesia 7–21 day old 7–21 day old
Given prenatally; 9 in females only Given postnatally 9
745 745 1117
239 239 425
277
472
(continued on next page)
2214
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 6 (continued) Peptide
Paradigm
ACTH/MSH frag. (4–10) ACTH/MSH frag. (4–10)
Benton visual retention test Cardiac orienting to novel stimuli Cardiac orienting to novel stimuli Classical conditioning Classical conditioning Conditioned taste aversion Digit symbol substitution test Discriminative learning
ACTH/MSH frag. (4–10) ACTH/MSH ACTH/MSH ACTH/MSH ACTH/MSH
frag. frag. frag. frag.
(4–10) (4–10) (4–10) (4–10)
ACTH/MSH frag. (4–10)
Phase
Species
Effect
Notes
Vol Page
Ref
Human Rabbit
Improvement Delayed
Elderly
Habituation
1 6
55 2S 97
359 228
Orienting
Rabbit
Enhanced
6
2S 97
228
Acquisition Extinction Retention
Rabbit Rabbit Rat Human
Worsening Faster Augmented version No change
3 3 1 1
715 715 207 55
478 478 501 359
Rat
Improvement
4
11
479
Rat
7
201
360
55 7
359 561
55 277
359 472
Discrimination stimulus Retention
Elderly In one out of ten rats
ACTH/MSH frag. (4–10)
Hunter delayed reaction apparatus
ACTH/MSH frag. (4–10) ACTH/MSH frag. (4–10)
Names and Faces test Passive avoidance response Retention
Human Rat
Enhanced at low doses inhibited at high doses No change Improvement
ACTH/MSH frag. (4–10) ACTH/MSH frag. (4–10)
Acquisition
Human Rat
No change Improvement
Extinction
Rat
Improvement
1
277
472
Reversal
Rat
No improvement
1
277
472
ACTH/MSH frag. (4–10) ACTH/MSH frag. (4–10) ACTH/MSH frag. (4–10)
Stroop test Thomson Box visual discrimination Thomson Box visual discrimination Thomson Box visual discrimination T-maze T-maze T-maze
Elderly 1 When admin. 3 prior to retention test, but not if admin. immediately after learning Elderly 1 1
Acquisition Extinction
Rat Rat Rat
7–21 day old 7–21 day old
6 6 10
2S 41 2S 41 1101
1 1 452
ACTH/MSH frag. (4–10) Actinomycin-D Amylin
Wechsler memory scale Passive avoidance Conditioned taste aversion
Elderly
Rat
1 13 13
55 927 961
359 308 79
Amylin Amylin
T-maze T-maze
Retention Retention
Human Rat No change Mouse Mouse
Improved No change Acted as a discrimination stimulus No change Impaired Intra-hypothalamic admin. Improvement Impairment
13 13
577 577
179 179
ANF, antiserum to ANF, antiserum to AT II
Active avoidance Extinction Passive avoidance response Consolidation Active avoidance
Rat Rat Rat
Facilitated Attenuated Improvement
13 13 21
957 957 407
46 46 37
AT II
Inhibitory shock avoidance
Retention
Rat
Impaired
16
1069
309
AT II AT II AT II
Passive avoidance Retention Passive avoidance behavior Passive avoidance behavior
Rat Rat Rat
Impaired Improvement No change
20 9 9
335 475 475
445 58 58
AT II AT II analog losartan
Plus maze Inhibitory shock avoidance
Rat Rat
Impaired Improved
20 16
335 1069
445 309
AT II analog saralasin BBS
Passive avoidance behavior Conditioned taste aversion Acquisition
Rat Rat
Improvement No change
Weak training ICV; strong training ICV ICV Hippocampal CA1 admin.; greater effect in right-side admin. Dentate gyrus admin. ICV ICV ICV with saralasin ICV Dentate gyrus admin.; AT IIinduced memory impairment ICV
9 2
475 61
58 301
ACTH/MSH frag. (4–10) ACTH/MSH frag. (4–10)
Retention
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2215
Table 6 (continued) Peptide
Paradigm
BBS
Species
Effect
Fixed interval responding
Rat
BBS BBS BBS Caerulein Caerulein
Non-reward operant task Reward operant task Taste aversion Active avoidance Morris water maze
Rat Rat Rat Rat Rat
Reduced responses after reinforcement Decreased Decreased No change Delayed Improvement
Caerulein
Morris water maze
Rat
Improvement
Caerulein
Morris water maze
Rat
Improved
Caerulein
Passive avoidance response Recall
Rat
Improved
Caerulein
Passive avoidance response Recall
Rat
Improved
Caerulein
Passive avoidance response Recall
Rat
Caerulein analog des-Gln-Caerulein Caerulein analog des-Gln-Caerulein Caerulein analog Leu-Nle-Caerulein Caerulein analog Leu-Nle-Caerulein Casein () analog D-TyrD-Phe3 CCK analog Boc-CCK-4 CCK analog Boc-CCK-4
Active avoidance response
Rat
Prevented ECSinduced amnesia Delayed
Extinction
Active avoidance response
Rat
Extinction
1435
55
4 4 3 10 13
1 1 61 843 1007
242 242 581 255 256
14
1073
535
12
699
537
11
1263
534
12
699
537
10
843
255
10
843
255
10
843
255
10
843
255
843
255
Amygdalakindled seizures
13
73
27
15 Central nucleus 19 of the amygdala admin.; post-trial injection 16 16
689 27
197 247
815 959
215 264
16 7 7 19 19 10 2 2 2 15
815 809 809 849 969 843 385 385 385 689
215 273 273 376 488 255 171 171 171 197
15
689
197
Rat Rat Rat
Increased Improved
Rat Rat
Rat Rat Rat Rat Human Rat Rat Rat Rat Rat
Impaired Potentiated amphetamine response Impaired No change Worsening No change Impaired recall Delayed Delayed Accelerated No change No effect
Rat
Increased
Extinction Acquisition Extinction Maintenance
12
10
Y-maze brightness discrimination Habituation Uphill avoidance task
Recall Acquisition Retention
Ref
ECS amnesia
Prevented ECSinduced amnesia Improved learning
Passive avoidance response Passive behavior response Passive behavior response Taste aversion conditioning Word recall Active avoidance response Conditioned feeding Conditioned feeding Conditioned feeding Conditioned place preference Habituation
Protein synthesis inhibitor-induced memory impairment Chronic ICV admin.; VIPinduced memory impairment Protein kinase-C inhibitor-induced memory impairment In rats with NMDA antagonistsinduced impairment. Protein kinase-C inhibitor-induced memory impairment ECS amnesia
Vol Page
ECS amnesia
Rat
Retention
Notes
Prevented ECSinduced amnesia Delayed
Passive avoidance response Recall
CCK CCK CCK CCK CCK CCK CCK CCK CCK CCK
CCK frag. CCK-8
Spatial memory
Rat
Active avoidance response Passive reward bar pressing
CCK-4 CCK-4 CCK-4 CCK-4 CCK-4 CCK-8 CCK-8 CCK-8 CCK-8 CCK-8
Extinction
Passive avoidance response Recall
CCK analog JMV-320 CCK analog L-365,260
frag. frag. frag. frag. frag. frag. frag. frag. frag. frag.
Phase
ICV ICV ICV, IV, or IP
(continued on next page)
2216
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 6 (continued) Peptide
Paradigm
CCK frag. CCK-8 CCK frag. CCK-8
Species
Effect
Non-reward operant task Passive avoidance response Recall
Rat Rat
CCK frag. CCK-8 CCK frag. CCK-8
Passive behavior response Passive behavior response
Acquisition Acquisition
Rat Rat
No change Prevented ECSinduced amnesia Improved Improved
CCK frag. CCK-8
Passive behavior response
Acquisition
Rat
Improved
CCK frag. CCK-8
Passive behavior response
Acquisition
Rat
Improved
CCK CCK CCK CCK CCK
Passive behavior response Retention Reward operant task Taste aversion conditioning Taste aversion conditioning Taste aversion conditioning Rat
Rat Rat Rat Rat Induced aversion Rat
Worsening Decreased Induced aversion ICV, IV, or IP Induced conditioning
frag. frag. frag. frag. frag.
CCK-8 CCK-8 CCK-8 CCK-8 CCK-8
Phase
CCK frag. CCK-8
Uphill avoidance task
CCK, antibody to
Discrimination task
CCK-8 CGRP, antiserum to CRF CRF
Free-operant avoidance Active avoidance response Active avoidance response Active avoidance response
CRF CRF
Arousal Conditioned taste aversion
CRF
CRF CRF binding protein
Fixed-interval food presentation Fixed-interval shock presentation Passive avoidance Shock-prod burying response Shock-prod burying test Active avoidance
CRF binding protein
Passive avoidance
Cycloheximide DDAVP Deltorphin
Passive avoidance Complex task execution Passive avoidance response Memory consolidation Fixed-interval food presentation Fixed-interval food presentation shock termination Delayed response in Working memory Wisconsin General Test apparatus Discrimination reversal Reversal Discrimination reversal Reversal task in Wisconsin General Test apparatus
CRF CRF CRF
Dermorphin Dermorphin
DSIP
DSIP DSIP
Retention
Notes
Vol Page
Ref
ECS amnesia
4 10
1 843
242 255
7 7
105 105
274 274
7
105
274
7
105
274
8 4 19 16 17
25 1 849 539 483
143 242 376 166 377
27
247
67
357
67 893 745 745
93 296 239 239
217 345
268 41
1199
384
ICV ICV; ECSinduced amnesia ICV; VPantiserum ECS-induced amnesia and VPantiserum
Central nucleus 19 of the amygdala admin.; post-trial injection Learning and recall Mouse Impairment Posterior 14 cingulate admin. Acquisition Rat Decreased avoidance 4 Extinction Rat Facilitated ICV 15 Acquisition and recall Mouse No change Given prenatally 9 Acquisition and recall Mouse Improvement in Given postnatally 9 females Monkey Increased ICV 4 Rat Induced aversion Third ventricle 20 admin. Monkey Increased responding ICV 10
Extinction
Improved
Monkey No change
ICV
10
1199
384
Rat Rat
Improved Decreased
ICV
13 13
927 1149
308 146
Rat Rat
Delayed Improvement
13 18
1149 711
146 227
Improved performance Rat Impaired Human Increased speed Mouse Increased step-down ICV latencies Monkey Increased responding ICV
18
711
227
13 12 11
927 1 591
308 76 421
10
1199
384
Monkey Decreased responding
10
1199
384
Monkey Worsening
2
1S 131 406
Monkey Worsening Monkey No change
2 2
1S 131 406 1S 131 406
Rat
ICV ICV; aged animals ICV
ICV
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2217
Table 6 (continued) Peptide
Paradigm
Dynorphin frag. (1–13)
Appetitive discrimination
Chicken Inhibited
Dynorphin frag. (1–13)
Avoidance conditioning
Chicken Inhibited
Endorphin ()
Act. behavior response
Recall
Rat
Endorphin ()
Acquisition
Endorphin ()
Fixed-ratio, fixed-interval food presentation Open field habituation
Monkey Decreased responding Rat Improved
Endorphin ()
Passive avoidance response Recall
Endorphin ()
Passive behavior response
Recall
Endorphin analog D-Ala2F5Phe4--endorphin
Delayed response in Wisconsin General Test apparatus Discrimination reversal task in Wisconsin General Test apparatus Discrimination reversal Conditioned aversion
Working memory
Reversed ECT8 induced amnesia but did not effect controls Rat Decreased avoidance 10 latency Rat Improved Reversed ECT8 induced- amnesia but did not affect controls Monkey Worsening 2
Reversal
Monkey No change
Reversal
Enkephalin analog D-Ala2- Delayed response in F5Phe4-Met-enkephalin Wisconsin General Test apparatus Enkephalin analog D-Ala2- Discrimination reversal F5Phe4-Met-enkephalin task in Wisconsin General Test apparatus Enkephalin analog D-Ala2-Met-enkephalinDelayed response in ethylamide Wisconsin General Test apparatus Enkephalin analog D-Ala2-Met-enkephalinDiscrimination reversal ethylamide task in Wisconsin General Test apparatus Enkephalin analog DPDPE Active avoidance response Enkephalin analog DPDPE Active avoidance response Enkephalin analog D-Phe4- Delayed response in Met-enkephalin Wisconsin General Test apparatus Enkephalin analog D-Phe4- Discrimination reversal Met-enkephalin task in Wisconsin General Test apparatus Enkephalin analog Fixed-ratio, fixed interval Metkephamide food presentation
Working memory
Monkey Worsening Rat Increased preference for conditioned stimulus, regardless of unconditioned stimulus Monkey Worsening
Reversal
Monkey No change
Working memory
Monkey Worsening
Reversal
Monkey No change
Acquisition Retention Working memory
Reversal
Endorphin analog D-Ala2F5Phe4--endorphin Enkephalin (D-Phe4-Met) Enkephalin (Met)
Phase
Species
Effect
Improved
Notes
Vol Page
Intermediate 14 medial hyperstriatum ventrale admin.; chicks Intermediate 14 medial hyperstriatum ventrale admin.; chicks Reversed ECT8 induced amnesia but did not affect controls 10
Ref
1165
95
1165
95
605
394
729
113
605
394
1117
425
605
394
1S 131 406
2
1S 131 406
2 In utero exposure 3
1S 131 406 881 520
2
1S 131 406
2
1S 131 406
2
1S 131 406
2
1S 131 406
Rat Impaired Rat Enhanced Monkey Worsening
13 13 2
885 340 885 340 1S 131 406
Monkey No change
2
1S 131 406
Rat
3
771
Decreased responding
305
(continued on next page)
2218
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 6 (continued) Peptide
Paradigm
Galanin Melanin
Species
Effect
Notes
Vol Page
Ref
Passive avoidance response Morris water maze Spatial memory
Mouse Rat
Decreased latency Improved
ICV In rats whose learning was inhibited by receiving MSG as neonates
16 12
1283 465
556 599
Melanin-concentrating hormone Melanin-concentrating hormone
Passive avoidance
Rat
Facilitated
15
757
343
Rat
Improved
20
1517
364
-(Tyr9) melanotropin(9–18) -(Tyr9) melanotropin(9–18) -(Tyr9) melanotropin(9–18) Melatonin
Active avoidance response
Extinction
Rat
Improved
Amygdala and hippocampus admin. ICV
7
11
540
T-maze
Reversal
Rat
Improved
ICV
7
11
540
T-maze
Spatial
Rat
Improved
ICV
7
11
540
Passive avoidance response Acquisition
Rat
Delayed
1
147
119
Melatonin
Passive avoidance response Extinction
Rat
Facilitated
Sham hypophysectomy but not hypophysectomy Sham hypophysectomy but not hypophysectomy
1
147
119
MIF-1
Delayed response in Wisconsin General Test apparatus Discrimination reversal task in Wisconsin General Test apparatus Lashly III maze Morris water maze
Working memory
Monkey Worsening
2
1S 131 406
Reversal
Monkey No change
2
1S 131 406
Learning Spatial memory
Mouse Rat
Improvement No change
11 12
527 465
Retention
Rat Rat
No change No change
2 8
1S 143 120 39 26
Retention
Rat
No change
8
39
26
Acquisition
Rat
Worsening
4
725
32
Reversal
Rat
No change
4
725
32
Extinction
Rat Rat Rat
Delayed No change No improvement
15 2 1
757 343 1S 143 120 277 472
Rat Improved Monkey Improved Rat Decreased errors
2 2 12
333 401 1S 131 406 929 450
MIF-1
MIF-1 MIF-1
MIF-1 MIF-1 analog Cyclo(Leu-Gly) MIF-1 analog Cyclo(Leu-Gly) MSH MSH MSH MSH MSH MSH MSH (␣) MSH (␣)
Phase
Extinction
Passive avoidance response
Passive avoidance response Conditioned avoidance response Conditioned avoidance response Fixed-ratio food presentation Fixed-ratio food presentation Passive avoidance Passive avoidance response Thomson Box visual discrimination Visual discrimination Discrimination reversal Visual discrimination learning
Acquisition
Reversal
Neonates In rats whose learning was inhibited by receiving MSG as neonates
Haloperidoltreated
Trend toward worsening
Medial anterior hypothalamic/ preoptic area admin.; no increase in responding to correct cues
115 599
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2219
Table 6 (continued) Peptide
Paradigm
Phase
MSH (␣)
Classically-conditioned fever response
MSH (␣)
Delayed response in Wisconsin General Test apparatus Discrimination reversal task in Wisconsin General Test apparatus Morris water maze
MSH (␣) MSH (␣)
MSH (␣)
Species
Effect
Notes
Vol Page
Ref
Classical conditioning Rat
11
1127
65
Working memory
Acted as unconditioned stimulus Monkey Improvement
2
1S 131 406
Reversal
Monkey No change
2
1S 131 406
Spatial memory
Rat
Improved
12
465
599
Acquisition
Rat
No change
4
145
222
Reversal
Rat
No change
4
145
222
Acquisition
Rat
Improved
4
145
222
Reversal
Rat
Improved
4
145
222
Acquisition
Rat
Facilitated
Hypophysectomy, 1 but not in controls Hypophysectomy, 1 but not control ECS amnesia 10
147
119
147
119
361
36
10 1
361 277
36 472
1
277
472
4 4 4 4 4 4 4 2
721 721 721 145 145 145 145 101
286 286 286 222 222 222 222 402
In rats whose learning was inhibited by receiving MSG as neonates
MSH (␣)
Operant auditory discrimination Operant auditory discrimination Operant visual discrimination Operant visual discrimination Passive avoidance response
MSH (␣)
Passive avoidance response Extinction
Rat
Delayed
MSH (␣)
Passive avoidance response Recall
Rat
MSH (␣) MSH (␣)
Passive avoidance response Thomson Box visual discrimination Thomson Box visual discrimination Y maze Y maze Y maze Y maze Y maze Y maze Y maze Y maze visual discrimination test Y maze visual discrimination test Y maze visual discrimination test Y maze
Recall Extinction
Rat Rat
Prevented ECSinduced amnesia No change No change
Reversal
Rat
Improvement
Acquisition Extinction Reversal Acquisition Acquisition Reversal Reversal Acquisition
Rat Rat Rat Rat Rat Rat Rat Rat
Improvement No change Improvement Improved No change Improved No change Improvement
Extinction
Rat
No change
2
101
402
Reversal
Rat
Improvement
2
101
402
Acquisition
Rat
No change
ICV
4
721
286
Y maze
Extinction
Rat
No change
ICV
4
721
286
Y maze
Reversal
Rat
Worsening
ICV
4
721
286
Passive avoidance response Recall
Rat
No change
ECS amnesia
10
361
36
Passive avoidance response Recall
Rat
Improved recall
With controls but 10 not ECS amnesia
361
36
MSH (␣) MSH (␣) MSH (␣)
MSH (␣) MSH MSH MSH MSH MSH MSH MSH MSH
(␣) (␣) (␣) (␣) (␣) (␣) (␣) (␣)
MSH (␣) MSH (␣) MSH analog [Nle,D-Phe]␣-MSH MSH analog [Nle,D-Phe]␣-MSH MSH analog [Nle,D-Phe]␣-MSH MSH analog Ac-Cys-GluHis-D-Phe-Arg-Trp-CysNH2 MSH analog Ac-Cys-GluHis-Phe-Arg-Trp-CysNH2
ICV ICV ICV
(continued on next page)
2220
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 6 (continued) Peptide
Paradigm
MSH analog Ac-Nle-AspHis-D-Phe-Arg-Trp-LysNH2 MSH analog Ac-Nle-GluHis-D-Phe-Arg-TrpDab-NH2 MSH analog Ac-Nle-GluHis-D-Phe-Arg-Trp-GlyLys-NH2 MSH analog Ac-Nle-GluHis-D-Phe-Arg-Trp-GlyLys-NH2 MSH analog Ac-Nle-GluHis-D-Phe-Arg-Trp-GlyLys-NH2 MSH analog Ac-Nle-GluHis-D-Phe-Arg-Trp-GlyLys-NH2 MSH analog Ac-Nle-GluHis-D-Phe-Arg-Trp-GlyNH2 MSH analog Ac-Nle-GluHis-D-Phe-Arg-Trp-GlyNH2 MSH analog Ac-Nle-GluHis-D-Phe-Arg-Trp-GlyOH MSH analog H-Cys-GluD-His-Phe-D-Arg-TrpD-Cys-OH MSH analog H-Cys-GluHis-Phe-D-Arg-Phe-DCys-OH MSH analog H-Cys-GluHis-Phe-D-Arg-Phe-DCys-OH MSH analog H-Cys-GluHis-Phe-D-Arg-Phe-DCys-OH MSH analog H-Cys-GluHis-Phe-D-Arg-Trp-CysOH MSH analog H-Cys-GluHis-Phe-D-Lys-Phe-CysOH MSH analog H-Cys-GluHis-Phe-D-Lys-Phe-DCys-OH MSH analog MK-771 MSH () MSH () MSH () MSH (␥) MSH (␥)
Phase
Species
Effect
Notes
Vol Page
Ref
Passive avoidance response Recall
Rat
No change
ECS amnesia
10
361
36
Passive avoidance response Recall
Rat
Prevented ECSinduced amnesia
ECS amnesia
10
361
36
Passive avoidance response Recall
Rat
Prevented ECSinduced amnesia
ECS amnesia
10
361
36
Thomson-Bryant discrimination test
Acquisition
Rat
No change
10
361
36
Thomson-Bryant discrimination test
Retention
Rat
No change
10
361
36
Thomson-Bryant discrimination test
Reversal
Rat
Impaired reversal
10
361
36
Passive avoidance response Recall
Rat
Prevented ECSinduced amnesia
10
361
36
Passive avoidance response Recall
Rat
No change
10
361
36
Passive avoidance response Recall
Rat
Prevented ECSinduced amnesia
ECS amnesia
10
361
36
Passive avoidance response Recall
Rat
Prevented ECSinduced amnesia
ECS amnesia
10
361
36
Thomson-Bryant discrimination test
Acquisition
Rat
No change
10
361
36
Thomson-Bryant discrimination test
Retention
Rat
No change
10
361
36
Thomson-Bryant discrimination test
Reversal
Rat
No change
10
361
36
Passive avoidance response Recall
Rat
No change
ECS amnesia
10
361
36
Passive avoidance response Recall
Rat
No change
ECS amnesia
10
361
36
Passive avoidance response Recall
Rat
No change
ECS amnesia
10
361
36
Fixed-ratio, fixed-interval shock avoidance Thomson Box visual discrimination Thomson Box visual discrimination Thomson Box visual discrimination Y maze visual discrimination test Y maze visual discrimination test
Monkey Increased
ECS amnesia
Acquisition
Rat
No Improvement
Under fixed-ratio 5 schedule 1
783
56
277
472
Extinction
Rat
No improvement
1
277
472
Reversal
Rat
Improvement
1
277
472
Acquisition
Rat
No change
2
101
402
Extinction
Rat
No change
2
101
402
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2221
Table 6 (continued) Peptide
Paradigm
Phase
Species
Effect
MSH (␥)
Reversal
Rat Rat
NPY
Y maze visual discrimination test Progressive-ratio operant schedule T-maze
Retention
Mouse
NPY
T-maze
Retention
Mouse
NPY frag. (20–36)
T-maze
Retention
Mouse
NPY frag. (20–36)
T-maze
Retention
Mouse
NPY frag. (26–36)
T-maze
Retention
Mouse
NT
Rewarded behavior
Rat
OT
Buschke restrictive reminding test Escape behavior
Human
Facial recognition test Memory comparison test One-trail step-through Retention paradigm Passive avoidance response Learning/recall Social discrimination response Social recognition
Human Human Rat
Stroop color-word test Y-maze brightness discrimination Passive avoidance T-maze T-maze
Recall
Human Rat
Retention Retention
Rat Mouse Mouse
Pancreastatin frag. (33–49) T-maze Retention Secretin Novel-object approach Secretin Passive avoidance Somatostatin Passive avoidance response
Mouse Rat Rat Rat
Somatostatin
Passive avoidance response Recall
Rat
Somatostatin (7–10) Somatostatin frag. (3–8) Somatostatin frag. (9–14) SP
Passive avoidance response Recall Passive avoidance response Recall Passive avoidance response Recall Morris water maze
Rat Rat Rat Rat
SP
Tunnel maze
Short and long term memory
Rat
SP SP analog [pGlu6]SP(6–11) SP frag. (1–7) TRH
Up-hill avoidance learning Up-hill avoidance learning
Retention Retention
Up-hill avoidance learning Active avoidance response
Retention
NPY
OT
OT OT OT OT OT OT OT OT PACAP Pancreastatin Pancreastatin
Vol Page
Ref
Worsening
2
101
402
Increased responding Perifornical region admin. Improvement ICV; weak training Impairment ICV; strong training Improvement ICV; weak training Impairment ICV; strong training No change ICV; weak training Decreased ICV responding Reduced initial storage Increased With central, but not peripheral, admin. No change No change No change
19
1667
60
10
963
177
10
963
177
10
963
177
10
963
177
10
963
177
16
1417
457
13
461
62
2
1
350
13 13 15
461 461 229
62 62 52
927 999
529 151
999
151
461 717
62 84
1067 1077 1077
536 180 180
1077 739 1067 293
180 83 536 568
1153
569
1153 1153 1153 275
569 569 569 504
85
385
Rat Rat
Prenatal exposure 11 Olfactory bulb 19 admin. Olfactory bulb 19 admin. No change 13 No change Newborns and 9 adults No change 14 Improvement 9 Improvement Scopolamine9 induced amnesia Improvement 9 Increased ICV 4 No change 14 Blocked ECTICV; ECT4 induced amnesia induced amnesia Decreased avoidance ICV 10 latency No change ICV 10 No change ICV 10 No change ICV 10 Improved Rats with lesions 17 performance of the hippocampus with or without fetal grafts Facilitated With pretrial, but 14 not post-trial, admin. Improvement 11 No change 11
163 163
224 224
Rat Rat
Improvement Improved
163 1309
224 152
Rat
Rat Rat Rat
Notes
No change Preserved recognition Increased
Rats with hypoxic amnesia
11 12
(continued on next page)
2222
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 6 (continued) Peptide
Paradigm
TRH
Active avoidance response
TRH
TRH analog MK-771
Fixed-ratio, fixed-interval food presentation Fixed-ratio, fixed-interval food presentation Fixed-ratio, fixed-interval food presentation Fixed-ratio, fixed-interval shock avoidance Passive avoidance response Radial arm maze Fixed-ratio, fixed-interval food presentation Fixed-ratio, fixed-interval food presentation Fixed-ratio, fixed-interval food presentation Radial maze (12-arm)
TRH analog RGH 2202
Active avoidance response
Recall
TRH analog RGH 2202
Active avoidance response
Recall
TRH analog RGH 2202 TRH analog RGH 2202 TSH
Passive avoidance response Recall Radial arm maze Spatial memory Fixed-interval food reward with fixed-ratio shock Fixed-interval food reward with fixed-ratio shock
TRH TRH TRH TRH TRH TRH analog MK-771 TRH analog MK-771 TRH analog MK-771
TSH
Tuftsin
Phase
Species
Effect
Rat
Monkey Recall Spatial memory
Working memory
Reversed age-related Aged animals impairment Monkey Decreased responding Pigeon Decreased responding Rabbit Decreased responding Increased Under fixed-ratio schedule Rat Improvement Rat Decreased errors Monkey Decreased responding Pigeon Decreased responding Rabbit Decreased responding Rat Improvement In medal septal lesioned but not controls Rat Reversed hypoxiaRats with induced amnesia hypoxic amnesia Rat Reversed age-related Aged animals impairment Rat Improvement Rat Decreased errors Pigeon Decreased When admin. alone Pigeon Increased When admin. with pentobarbital, chlordiazepoxide, or alcohol Monkey Increased responding ICV
Urocortin
Fixed-interval food presentation Fixed-interval shock presentation Conditioned taste aversion
Vasotocin
Passive avoidance behavior Retention
Chicken Increased
Vasotocin
Acquisition
Rat
No change
Reversal
Rat
Improvement
VIP
Thompson-Bryant discrimination box Thompson-Bryant discrimination box Morris water maze
Rat
Impairment
VIP VIP VIP
Passive avoidance Passive avoidance T-maze
Rat Rat Mouse
Impairment No change Impairment
VP VP
Active behavior response Cognition
Rat Human
Slowed Improved
Tuftsin
Vasotocin
Notes
Monkey Decreased responding Rat Induced aversion
Retention
Extinction Activation, alertness, and fatigue
ICV
Vol Page
Ref
12
1309
152
4
177
23
4
177
23
4
177
23
5
783
56
12 12 4
1309 1309 177
152 152 23
4
177
23
4
177
23
10
121
240
12
1309
152
12
1309
152
12 12 5
1309 1309 809
152 152 591
5
809
591
10
1199
384
10
1199
384
345
41
1221
122
383
35
6
383
35
14
1073
535
14 14 11
1067 1067 933
536 536 176
7 12
213 1385
290 429
Third ventricle 20 admin. ICV or IP; 2 day 5 old chicks 6
Chronic ICV admin. Central admin. Peripheral admin. ICV; effect antagonized by arecoline, naloxone and ST 587, but not CCK or NPY ICV
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2223
Table 6 (continued) Peptide
Paradigm
Phase
Species
Effect
Notes
Vol Page
Ref
VP
Conditioned aversion response
Acquisition
Rat
ICV
7
213
290
VP
Escape behavior
Rat
Did not act as unconditioned stimulus Increased
2
1
350
VP
Habituation
Mouse
Increased
With central, but not peripheral, admin. Transgenic admin.
16
1329
362
VP VP
Recall Recall
Human Human
Enhanced free recall No change
12 12
1379 1379
391 391
Retention
Rat
Impairment
15
229
52
Human
No change
12
1393
20
VP
Incidental memory task Oddball paradigm word recall One-trail step-through paradigm Orienting response and stimulus mismatch Passive avoidance
Mouse
No change
1329
362
VP VP
Passive avoidance behavior Recall Passive avoidance response Learning/recall
Rat Rat
493 927
550 529
VP VP VP
Passive avoidance response Retention Passive behavior response Acquisition Passive behavior response Acquisition
Rat Rat Rat
Impairment Increased step-down latency (improvement) in females Improvement No change Improved
Transgenic 16 admin. Given prenatally 8 Prenatal exposure 11
11 7 7
633 105 105
317 274 274
VP VP
Radial maze (12 arm) Social discrimination response Social recognition
Working
Rat Rat
1 19
261 999
66 151
19
999
151
Thompson-Bryant discrimination box Thompson-Bryant discrimination box Thompson-Bryant discrimination test Thompson-Bryant discrimination test Thompson-Bryant discrimination test Thomson-Bryant discrimination test Thomson-Bryant discrimination test Conditioned avoidance for smoking cessation Conditioned avoidance for smoking cessation Emotionally-influenced free recall Impaired free recall performance Radial maze (24-arm)
VP VP
VP VP VP VP VP VP VP VP VP (Lys) VP (Lys) VP (Lys) VP (Lys) VP analog 8-D-arginineVP VP analog 8-L-argininedeamino-VP VP analog DAVP VP analog DDAVP
Neonatal exposure
ICV ICV; ECSinduced amnesia
Rat
No change Preserved recognition Increased
Acquisition
Rat
No change
6
383
35
Reversal
Rat
Improvement
6
383
35
Acquisition
Rat
No change
Given prenatally
8
493
550
Reversal
Rat
No change
Given prenatally
8
493
550
Reversal recall
Rat
Impairment
Given prenatally
8
493
550
Acquisition
Mouse
No change
10
237
549
Reversal
Mouse
Improvement
237
549
Acquisition
Human
Improvement
In albino, but not 10 hooded, rats 3
527
164
Extinction
Human
Accelerated
3
527
164
Human
No change
Twins
6
297
170
Human
No change
Twins
6
297
170
Working memory
Rat
No change
3
725
67
Radial maze (24-arm)
Working memory
Rat
No change
3
725
67
Radial maze (12-arm) Active avoidance
Working Acquisition
Rat Rat
No change Enhanced at low dose and decreased at high dose
1 12
261 471
66 493
Olfactory bulb admin. Olfactory bulb admin.
(continued on next page)
2224
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 6 (continued) Peptide
Paradigm
Phase
Species
Effect
Notes
Vol Page
Ref
VP analog DDAVP
Active avoidance
Extinction
Rat
Inhibition
Dose-dependent effect
12
471
493
VP VP VP VP
Active avoidance test Active avoidance test Benton visual retention test Color and shape discrimination Delayed word list recall Immediate recall of sentences Immediate word list recall Implicational sentence recall
Acquisition Extinction
Rat Rat Human Human
Improved Delayed No change Improvement
6 6 3 3
23 23 627 627
221 221 29 29
Recall
Human Human
No change Improved
11 6
1313 397
31 548
Recall Recall
Human Human
11 11
1313 473
31 33
11 11 7 7
1313 633 563 563
31 317 172 172
7 7
563 563
172 172
16 1 5
1327 261 819
30 66 34
4
707
28
analog analog analog analog
DDAVP DDAVP DDAVP DDAVP
VP analog DDAVP VP analog DDAVP VP analog DDAVP VP analog DDAVP
Males Males
Paper folding test Passive avoidance response Retention Passive behavior response Acquisition Passive behavior response Acquisition
Human Rat Rat Rat
Impaired Dose-dependent impairment or facilitation No change Improvement No change Improved
VP analog DDAVP VP analog DDAVP
Passive behavior response Passive behavior response
Acquisition Acquisition
Rat Rat
Improved Improved
VP analog DDAVP VP analog DDAVP VP analog DDAVP
Phrase recall Radial maze (12-arm) Sentence comprehension and recall
Working
Human Rat Human
No change Worsening Improvement
VP analog DDAVP
Human
Improved attention
Acquisition
Human Rat
No change Improvement
11 3
1313 521
31 100
Extinction
Rat
No change
3
521
100
Reversal
Rat
Impairment
3
521
100
VP analog DDAVP
Sternberg item recognition task Stroop color word test Thomson Box visual discrimination Thomson Box visual discrimination Thomson Box visual discrimination Word list recall
Human
9
1361
430
VP analog DDAVP
Word recall
Human
Improved recent recall, worsened distant recall Improved
16
179
63
VP analog DDAVP
Y-maze brightness discrimination Y-maze brightness discrimination Classical conditioning
Acquisition
Rat
Improvement
9
717
84
Recall
Rat
Improvement
9
717
84
Acquisition
Rabbit
No change
4
37
229
VP analog Deaminodicarba-arginine-8-VP
Classical conditioning
Acquisition
Rabbit
No change
4
37
229
VP analog Deaminodicarba-arginine-8-VP
Classical conditioning
Extinction
Rabbit
Accelerated
4
37
229
VP analog Deaminodicarba-arginine-8-VP
Classical conditioning
Extinction
Rabbit
No change
4
37
229
VP VP VP VP
analog analog analog analog
DDAVP DDAVP DDAVP DDAVP
VP analog DDAVP VP analog DDAVP VP analog DDAVP VP analog DDAVP
VP analog DDAVP VP analog Deaminodicarba-arginine-8-VP
Phase-shiftinduced amnesia Phase-shiftinduced amnesia
Effect seen in males but not females Males
In men but not women Newborns but not adults Newborns and adults Classicallyconditioned bradycardia Classicallyconditioned nictitating membrane response Classicallyconditioned bradycardia Classicallyconditioned nictitating membrane response
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2225
Table 6 (continued) Peptide
Paradigm
Phase
Species
Effect
VP analog desGlyNH2DDAVP VP analog desGly-NH2VP VP analog DGAVP VP analog DGAVP
Radial maze (24-arm)
Working memory
Rat
Radial maze (12-arm)
Working
Active behavior response Active behavior response
Extinction Extinction
VP analog DGAVP
Buschke restrictive reminding test
VP analog DGAVP
Cardiac orienting to novel stimuli Cardiac orienting to novel stimuli Facial recognition test Memory comparison test
VP analog DGAVP VP analog DGAVP VP analog DGAVP
Notes
Vol Page
Ref
Improvement
3
725
67
Rat
No change
1
261
66
Rat Rat
Slowed Slowed
7 7
563 563
172 172
Human
13
461
62
6
2S 97
228
Phase-shiftinduced amnesia
Habituation
Rabbit
Increased word recall and delayed recognition Delayed
Orienting
Rabbit
Enhanced
6
2S 97
228
Human Human
13 13
461 461
62 62
13
461
62
7 11 11
213 633 633
290 317 317
Stroop color-word test
Human
No change Diminished reaction time No change
Active behavior response Extinction Passive avoidance response Retention Passive avoidance response Retention
Rat Rat Rat
Faster Improvement Improvement
Thompson-Bryant Acquisition discrimination box Thompson-Bryant Reversal discrimination box Passive avoidance response Retention
Rat
No change
6
383
35
Rat
No change
6
383
35
Rat
Improvement
11
633
317
Passive avoidance response Retention
Rat
Improvement
11
633
317
Passive avoidance response Retention Passive avoidance behavior Retention
Rat No change Chicken Enhanced avoidance
11 4
633 401
317 123
Passive avoidance response Post-learning application
Rat
No change
6
2S 177 566
VP, antibody to
Passive avoidance response Retention
Rat
Impairment
6
2S 177 566
VP, antibody to VP, antibody to
Passive avoidance response Retention Passive avoidance response Retention
Rat Mouse
Impairment Retention deficit
6 4
2S 177 566 265 306
VP analog DGAVP VP analog dPTyr(me) ArgVP VP analog D-ZNC(C)PR VP analog D-ZNC (C-OMe)PR VP analog Pressinoic acid VP analog Pressinoic acid VP analog ZNC(C)PR (VP 4–8) VP analog ZNC(COMe)PR VP analog ZNCPR dimer VP frag. L-prolyl-Larginyl-glycinamide VP, antibody to
ICV; VP treated
ICV but not IP; 2 day old chicks Dorsal septum admin. after learning Hippocampal admin. ICV ICV
2226
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 7 Stereotyped and other behaviors sorted by peptide Peptide
Behavior
Species
Effect
ACTH ACTH ACTH ACTH
Distress vocalization Grooming Grooming Grooming
Chicken Rat Rat Rat
Increased Increased Increased Increased
ACTH ACTH
Grooming Grooming
Rat Rat
Increased Increased
ACTH
Grooming
Rat
Increased
ACTH
Grooming
Rat and mouse
Increased
ACTH
Mouse
No change
ACTH ACTH
Myoclonus (serotonergic behavior) Vomiting Yawning
Dog Rat
No change Increased
ACTH frag. (1–24) ACTH frag. (1–24)
Anxiety behaviors Grooming
Chicken Rat
Increased Increased
ACTH frag. (1–4)
Grooming
Rat
Enhanced
ACTH frag. (4–10) ACTH frag. (4–10) ACTH/MSH 4–10 analog ORG 2766 ACTH/MSH frag. analogs, C-frag. elongated or cyclized Adrenal Peptide E (Frog)
Anxiety behaviors Distress vocalization Rotation
Chicken Chicken Rat
Increased Increased Decreased
Grooming
Rat
Increased
Mouse
No change
Rat Dog Rat
BBS BBS BBS
Morphine-induced behaviors Grooming Vomiting Amphetamine-induced stereotypy Apomorphine-induced stereotypy Bite/scratch behavior Grooming Grooming
Vol
Page
Ref
11 7 4 5
915 597 907 713
414 476 202 546
6 7
369 651
562 160
16
1263
434
8
841
159
10
5
439
6 16
1S 173 1263
597 434
11 4
915 833
414 6
7
1
134
11 11 14
915 915 1317
414 414 10
5
1197
232
ICV
17
1291
97
No change No change Increased
ICV ICV ICV
17 6 9
589 1S 173 475
91 597 58
Rat
Increased
ICV
9
475
58
Mouse Rat Rat
Increased Increased Increased
7 2 8
835 2S 179 237
47 198 303
Grooming Grooming Grooming
Rat Rat Rat
Increased Increased Increased
12 13 4
761 1215 693
182 99 355
BBS
Grooming
Rat
Increased
7
557
352
BBS
Grooming
Rat
Increased
9
1S 245
260
BBS
Grooming
Rat
Increased
9
1S 245
260
BBS
Grooming
Rat
Increased
Intraspinal admin. ICV With admin. to multiple brain areas ICV ICV; developed over 8 days ICV; effect antagonized by neuroleptics ICV; antagonized by morphine With admin. to nucleus tractus solitarius but not nucleus accumbens With admin. to nucleus tractus solitarius but not nucleus accumbens With ICV but not IV admin.; no antagonism by haloperidol, morphine, naloxone, or NT. No effect of hypophysectomy or adrenalectomy
BBS BBS BBS
4
907
202
Amylin AT AT II AT II
Notes ICV; blocked by naloxone ICV Pre-optic, anterior hypothalamic admin. ICV ICV; area of action localized to anterior-ventral third ventricle ICV; not antagonized by nitric oxide synthase inhibitor ICV; greater effect in rats than mice 5-hydroxytryptophaninduced ICV ICV; antagonized by nitric oxide synthase inhibitor ICV; effect antagonized by naloxone In MSH-induced grooming; no effect alone
Nigrostriatal lesions and amphetamine-induced turning More potent than unmodified frag.
(continued on next page)
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2227
Table 7 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
BBS
Grooming
Rat
Increased
6
369
562
BBS
Grooming
Rat
Increased
6
1179
564
BBS
Grooming
Rat
Increased
6
1179
563
BBS BBS BBS BBS BBS
Grooming and scratching Licking Licking Respiration Righting
Rat Rat Rat Rat Rat
Increased Decreased Decreased Increased Impaired
ICV; different doses induce different types of grooming Effect enhanced by haloperidol, antagonized by naloxone and NT ICV; antagonized by haloperidol Neonates
10 17 16 4 2
529 107 903 277 1S 99
259 175 523 395 187
BBS
Scratching
Mouse
Increased
9
909
567
BBS
Scratching
Rat
Increased
6
1179
563
BBS BBS Bradykinin Calcitonin Calcitonin
Sighing Vomiting Vomiting Respiration Writhing
Rat Dog Dog Rat Mouse
Increased No change No change Increased Increased analgesia
4 6 6 4 6
277 1S 173 1S 173 277 3S 277
395 597 597 395 200
Casomorphin analog BCH 325 Casomorphin analog BCH 325
Jerking
Rat
Decreased
16
635
447
Yawning
Rat
Normalized
16
635
447
Casomorphin analogs
Yawning
Rat
Decreased
13
69
464
Casomorphin frag. (1–5) CCK CCK CCK
Distress vocalization Anxiety behavior Respiration Stretching
Chicken Rat Cat Mouse
Decreased Increased Increased tidal volume Decreased
5 16 9 10
829 1313 809 31
415 263 246 9
CCK analog Boc-CCK-4
Anxiety behavior
Rat
No change
19
27
247
CCK frag. CCK-8
Anxiety behavior
Rat
No change
19
27
247
CCK-4 CCK-4 CCK-8
Circling Defecation Catalepsy
Rat Rat Rat
No change No change Blocked
7 6 6
809 91 237
367 275 253
CCK-8 CCK-8 CCK-8
Circling Defecation Rotation
Rat Rat Rat
Increased No change Increased
7 6 18
809 91 1161
367 275 396
CCK-8 CCK-8, desulfated Ceruletide
Rumination Circling Catalepsy
Sheep Rat Rat
No change No change Decreased
5 7 10
81 809 779
238 367 250
Ceruletide analogs Ceruletide analogs
Catalepsy Convulsions
Mouse Mouse
Increased Decreased
3 3
701 701
610 610
ICV ICV; alcohol-induced righting impairment Intrathecal admin.; not antagonized by D-Pro2,DTrp7,9-SP and phenoxybenzamine ICV; antagonized by naloxone ICV ICV ICV ICV ICV, IP, and IV, but no effect SC; Ach-induced writhing Bromocriptine-induced jerks Bromocriptine-evoked yawning that was inhibited by reserpine ICV; apomorphine-induced yawning ICV; chicks Ventral medulla admin. ICV; abdominal irritantinduced stretching Central nucleus of the amygdala admin. Central nucleus of the amygdala admin. ICV Nucleus accumbens admin. ICV; endorphin-induced catalepsy; effect blocked by frontal cortex lesion ICV Nucleus accumbens admin. Animals pre-treated with peptide prior to nigrostriatal lesions; dopaminedepletion-induced turning ICV Haloperidol-induced catalepsy Hexobarbital-induced sleep Harman-induced convulsions
(continued on next page)
2228
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 7 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
Ceruletide analogs
Gnawing
Mouse
Increased
Methylphen-induced gnawing
2
2S 65
609
Ceruletide analogs CGRP
Ptosis Grooming
Mouse Rat
Increased Increased
2 17
2S 65 1183
609 294
CGRP CRF
Grooming Boxing behavior
Rat Rat
Decreased Decreased
5 6
861 891
297 107
CRF CRF CRF
Grooming Grooming Grooming
Rat Rat Rat
Increased Increased Increased
13 6 20
1149 3S 53 509
146 392 571
CRF
Grooming
Rat and mouse
Increased
8
841
159
CRF CRF antagonist ␣-helical CRF Cyclo (Leu-Gly)
Huddling and lying down Freezing behavior
Monkey Rat
Increased Decreased
4 15
217 1303
268 237
Rat
Decreased
8
39
26
Cyclo (Leu-Gly)
Apomorphine-induced stereotypy Rotation
Rat
No change
8
39
26
Dermorphin
Catalepsy
Rat
Increased
6
3S 165
59
Dermorphin
Catalepsy
Rat
Decreased
6
3S 165
59
Dynorphin analog desTyr1-dynorphin (1–13) Dynorphin frag (1–13) Dynorphin frag. (1–13)
Grooming
Rat
Increased
4
833
6
Barrel rolling Grooming
Rat Rat
Increased Increased
8 4
1089 833
201 6
Dynorphin frag. (1–13)
Grooming
Rat and mouse
Increased
1
341
578
Dynorphin Dynorphin Dynorphin Dynorphin Dynorphin
Hind limb paralysis Shaking Circling Circling Circling
Rat Rat Rat Rat Rat
Increased Increased Increased Increased Increased
4 8 8 8 8
631 1089 837 837 837
168 201 186 186 186
Dynorphin frag. (2–17) Endorphin (␣)
Circling Distress vocalization
Rat Chicken
Increased Decreased
8 5
837 823
186 573
Endorphin ()
Catalepsy
Rat
Increased
6
237
253
Endorphin ()
Distress vocalization
Chicken
Decreased
5
823
573
Endorphin ()
Righting
Rat
Impaired
2
1S 99
187
Endorphin () Endorphin ()
Startle Trembling
Rat Rat
Decreased Decreased
2 2
1S 137 1S 99
234 187
Endorphin (␥)
Amphetamine-induced stereotypy
Rat
Decreased
2
9
285
Endorphin (␣)
Distress vocalization
Chicken
Decreased
5
823
573
Enkephalin
Vocalization
Sheep
Decreased
9
1221
417
frag. frag. frag. frag. frag.
(1–13) (1–13) (1–6) (1–7) (1–8)
Antagonized by nitric oxide synthase inhibitor ICV Medial frontal cortex admin.; carbachol-induced boxing behavior ICV ICV but not SC ICV; not antagonized by melanocortin-4 receptor antagonist ICV; greater effect in rats than mice; not antagonized by dexamethasone ICV Foot-shock-induced behavior Rats treated chronically with haloperidol Apomorphine-induced rotation in rats with unilateral nigrostriatal lesions ICV; effects increased by caudate lesion ICV; in dermorphin-tolerant but not morphine-tolerant rats ICV; effect antagonized by naloxone ICV or IV ICV; effect antagonized by naloxone ICV; not reversible by naloxone Intrathecal admin. ICV or IV Substantia nigra admin. Substantia nigra admin. Substantia nigra admin.; not antagonized by naloxone Substantia nigra admin. With ICV, but not IP, admin. ICV; effects enhanced by frontal cortex ablation With ICV, but not IP, admin.; chicks ICV; alcohol-induced righting impairment ICV; alcohol-induced tremors Effect antagonized by 2bromo-ergocryptine and naloxone With ICV, but not IP, admin. ICV
(continued on next page)
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2229
Table 7 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
Enkephalin (Leu) Enkephalin (Met) Enkephalin analog DADLE
Circling Shaking Respiration
Rat Fish Rat
Increased Increased Increased
Substantia nigra admin.
8 6 3
837 139 1031
186 86 225
Enkephalin analog DADLE Enkephalin analog DAGO
Respiration Respiration
Rat Rat
3 3
1023 1031
236 225
Enkephalin analog DAMA Enkephalin analog FK33– 824 Galanin Galanin
Shaking Respiration
Rat Mouse
Increased Increased at low dose decreased at high dose Increased Decreased rate
4 9
239 777
155 378
Anxiety behavior Freezing behavior
Mouse Rat
No change No change
16 15
1283 1303
556 237
Galanin
Grooming
Rat
No change
11
1001
304
Galanin
Rat
No change
15
1303
237
Gastrin (1–17) Gastrin-releasing peptide Gastrin-releasing peptide
Opiate withdrawal behavior Rumination Bite/scratch behavior Licking
Sheep Mouse Rat
Increased Increased Decreased
5 7 17
81 835 107
238 47 175
Gastrin-releasing peptide Kassinin L-pyroglutamyl-L-leucine
Licking Bite/scratch behavior Jumping
Rat Mouse Rat
Decreased Increased Delayed
16 7 4
903 835 417
523 47 295
MCH
Anxiety behavior
Rat
Decreased
17
171
207
Melatonin MIF-1
Startle response Catalepsy
Rat Mouse
Decreased Prevented tolerance
2 8
1S 155 1051
118 380
MIF-1
Catalepsy
Rat
Decreased
2
105
85
MIF-1
Circling
Rat
No change
2
189
148
MIF-1 MIF-1
Defecation Head turning
Rat Rat
Decreased No change
2 2
1S 143 189
120 148
MIF-1 MIF-1
Head twitch Oral act. (serotonergic behavior)
Mouse Rat
No change No change
2 14
189 1159
148 206
MIF-1 MIF-1 analog 3(R)-[(2(S)pyrrolidinylcarbonyl) amino]-2-oxo-1pyrrolidineacetamide MIF-1 analog Cyclo(Leu-Gly) MIF-1 analog parepide
Depressive behaviors Rotation
Human Rat
Decreased Increased
4 18
297 1209
559 363
Sniffing
Rat
Increased
5
7
310
Catalepsy
Rat
Increased
5
883
381
MIF-1 analog YPLG
Catalepsy
Mouse
No change
8
1051
380
MSH
Defecation
Rat
Increased
1
147
119
MSH MSH MSH
Defecation Grooming Vocalization
Rat Rat Chicken
Increased Increased Decreased
2 2 11
1S 143 333 647
120 401 413
MSH (␣)
Anxiety behavior
Rat
Increased
17
171
207
Nucleus tractus solitarius admin. Third ventricle admin. Nucleus tractus solitarius admin. ICV
ICV Foot-shock-induced behavior Paraventricular nucleus admin.
ICV Intraspinal admin. Admin. simultaneously with neuromedin B-10 Intraspinal admin. Morphine-withdrawalinduced jumping Admin. to medial preoptic area Haloperidol-induced catalepsy Haloperidol-induced catalepsy ICV; dopamine- or 5HTstimulated circling Novelty-induced defecation Striatally-evoked head turning 5HT-stimulated twitch 6-hydroxydopaminelesioned rats treated with desipramine Apomorphine- or L-DOPAinduced rotation in 6hydroxydopamine-lesioned rats Apomorphine-induced sniffing Haloperidol-induced catalepsy Haloperidol-induced catalepsy Passive-avoidance-induced defecation Novelty-induced defecation ICV Separation-induced vocalization in chicks Admin. to medial preoptic area
(continued on next page)
2230
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 7 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
MSH (␣) MSH (␣) MSH (␣)
Grooming Grooming Grooming
Rat Rat Rat
Increased Increased Increased
2 4 16
101 721 821
402 286 110
MSH (␣)
Grooming
Rat
Increased
18
393
470
MSH (␣)
Grooming
Rat
Increased
7
1
134
MSH (␣)
Grooming
Rat
Increased
Not antagonized by ␥-MSH ICV ICV; antagonized by diazepam or baclofen ICV; antagonized by melanin-concentrating hormone Enhanced by ACTH frag. (1–4) Ventral tegmental area admin.; effect enhanced by bicuculline, blocked by atropine
12
203
127
MSH (␥) MSH analog [Nle4, DPhe7]-␣ MSH MSH analogs MSH antagonist [D-Trp7, Ala8, D-Phe10) ␣-MSH(6–11)amide Neuromedin B Neuromedin B-10
Grooming Grooming
Rat Rat
No change Increased
2 4
101 721
402 286
Grooming Walking pattern
Rat Rat
Increased Increased recovery
6 16
1185 319
505 433
Bite/scratch behavior Licking
Mouse Rat
Increased Decreased
7 17
835 107
47 175
NP-E-I NPY NPY
Grooming Anxiety behavior Anxiety behaviors
Rat Rat Mouse
18 14 19
393 909 359
470 428 387
NPY NPY NPY antibody
Catalepsy Grasping Barrel rolling
Rat Rat Mouse
Increased Decreased Decreased at low dose, increased at high dose Increased Increased Increased
15 15 15
799 799 607
54 54 579
NPY antibody
Circling
Mouse
Increased
15
607
579
NPY frag. (1–30) NPY frag. (1–30) NT NT NT NT
Catalepsy Grasping Grooming Respiration Respiration Righting
Rat Rat Rat Monkey Rat Rat
No effect No effect No change Decreased No change Impaired
15 15 4 5 4 2
799 799 493 125 277 1S 99
54 54 511 366 395 187
NT NT NT
Rotation Vomiting Yawning
Rat Dog Rat
Increased Increased Decreased
6 6 12
1057 1S 173 755
389 597 398
NT analog [D-Trp11]NT
Yawning
Rat
Decreased
12
755
398
OT OT OT
Barrel rolling Grooming Grooming
Rat Rat Rat
Increased Increased Increased
6 12 9
747 113 1S 223
492 405 522
OT OT
Grooming Grooming and scratching
Rat Rat
Increased
18 2
1389 1
153 350
ICV
Admin. after sciatic nerve crush Intraspinal admin. Admin. simultaneously with gastrin-releasing peptide ICV; antagonized by MSH ICV ICV; anxiolytic effect via Y1 receptors and anxiogenic via Y2 receptors ICV ICV With admin. to ventrolateral thalamus but not other brain areas With admin. to ventromedial thalamus but not other brain areas ICV ICV PVN and IV admin. ICV ICV ICV; alcohol-induced righting impairment Substantia nigra admin. ICV ICV; apomorphine-induced yawning ICV; apomorphine-induced yawning ICV ICV Ventral tegmental area admin.; effect not altered by sex, sex-steroids or dopamine antagonists ICV With central, but not peripheral, admin.; in nonstressful, but not stressful, conditions
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2231
Table 7 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
OT
Human
Decreased
In a single patient
13
1083
139
Rat Rat Dog Rat Rat Mouse Rat
Increased Increased Increased Increased No change Increased Increased
Intrathecal admin. ICV ICV ICV Intraspinal admin. ICV
6 6 6 10 20 7 9
567 1191 1S 173 559 1445 835 475
284 538 597 11 14 47 58
Rat
Increased
ICV
9
475
58
Sauvagine
Obsessive compulsive behavior Startle response Tail movement Vomiting Yawning Anxiety behavior Bite/scratch behavior Amphetamine-induced stereotypy Apomorphine-induced stereotypy Grooming
Rat
Increased
6
3S 53
392
Sauvagine Secretin Secretin Secretin Somatostatin Somatostatin
Grooming Defecation Gnawing Grooming Barrel rolling Scratching
Rat Rat Rat Rat Rat Mouse
Decreased Increased No change No change Increased Increased
6 4 4 4 10 9
3S 53 739 739 739 1153 909
392 83 83 83 569 567
Somatostatin frag. (3–8), (9–14), or (7–10) SP SP SP
Barrel rolling
Rat
No change
ICV but not SC; in fasted, but not satiated animals SC but not ICV ICV ICV ICV ICV Intrathecal admin.; not antagonized by D-Pro2,DTrp7,9-SP and phenoxybenzamine ICV
10
1153
569
Bite/scratch behavior Biting Boxing behavior
Mouse Mouse Rat
Increased Increased Increased
Intraspinal admin. Spinal subarachnoid admin. Medial frontal cortex admin.; carbachol-induced boxing behavior
7 20 6
835 301 891
47 469 107
SP SP
Chafing movements Grooming
Fish Mouse
Increased Increased
6 6
139 363
86 217
SP SP
Grooming Grooming
Rat Rat
Increased Increased
1 7
103 557
262 352
SP SP SP SP
Licking Respiration Rigidity Risk assessment behavior
Mouse Rat Rat Rat
Increased No change Reversed Increased
20 4 1 20
301 277 103 1437
469 395 262 126
SP
Rotation
Rat
Normalized
18
1161
396
SP SP
Scratching Scratching
Mouse Mouse
Increased Increased
4 6
517 363
249 217
SP
Scratching
Mouse
Increased
9
909
567
SP SP SP SP antagonist [D-Pro2, D-Phe7, D-Trp9]-SP
Scratching Scratching and licking Vomiting Boxing behavior
Mouse Rat Dog Rat
Increased Increased No change Decreased
20 3 6 9
301 49 1S 173 1S 117
469 485 597 521
OT OT OT OT Pancreatic polypeptide Ranatensin Saralasin Saralasin
ICV; different effectiveness in different strains and ages; enhanced by naloxone but not phenoxybenzamine or haloperidol ICV ICV; not antagonized by morphine Spinal subarachnoid admin. ICV ICV; in haloperidol treated Dorsal periaqueductal gray matter admin. Animals pre-treated with peptide prior to nigrostriatal lesions; dopaminedepletion-induced turning Intrathecal admin. ICV; different effectiveness in different strains and ages Intrathecal admin.; antagonized by D-Pro2,DTrp7,9-SP and phenoxybenzamine Spinal subarachnoid admin. Intrathecal admin. ICV Carbachol-induced boxing behavior
(continued on next page)
2232
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 7 (continued) Peptide 2
Behavior
Species
Effect
Notes
Vol
Page
Ref
Carbachol-induced boxing behavior Carbachol-induced boxing behavior
9
1S 117
521
9
1S 117
521
4 20
763 1437
219 126
4
763
219
SP antagonist [D-Pro , D-Trp7,9]-SP SP antisera
Boxing behavior
Rat
Decreased
Boxing behavior
Rat
No change
SP frag. (1–7) SP frag. (7–11)
Grooming Risk assessment behavior
Mouse Rat
Decreased Increased
SP frag. analog pyroglutamyl-SP (7–11) Substance K Tachykinin agonist GR73632 Tachykinin agonist GR73632 Tachykinin agonist GR73632 Tachykinin antagonist [Sar9,Met(O2)11]SP TK PG-KII TRH
Grooming
Mouse
Increased
Bite/scratch behavior Biting
Mouse Mouse
Increased Increased
Intraspinal admin. Spinal subarachnoid admin.
7 20
835 301
47 469
Licking
Mouse
Increased
Spinal subarachnoid admin.
20
301
469
Scratching
Mouse
Increased
Spinal subarachnoid admin.
20
301
469
Grooming
Rat
Increased
18
1349
87
Grooming Grooming
Rat Rat
Increased Increased
18 9
825 283
437 565
TRH
Licking
Rat
Increased
2
1S 99
187
TRH
Respiration
Rat
Increased
4
277
395
TRH
Righting
Rat
Improved
2
1S 99
187
TRH TRH TRH analog RGH 2202 TRH analog RX 77386 TRH analog RX 77386 Urotensin I
Shaking Tail movement Grooming Grooming Shaking Grooming
Rat Rat Rat Rat Rat Rat
Increased Increased Increased Increased Increased Increased
Nucleus basalis magnocellularis admin. ICV ICV; effect antagonized by haloperidol, naloxone and NT ICV; punished licking in water-deprived, ethanoltreated rats ICV; decreased tidal volume ICV; alcohol-induced righting impairment ICV Intrathecal admin.
4 6 12 11 11 6
239 1191 1309 897 897 3S 53
155 538 152 258 258 392
Urotensin I Vasotocin VIP antagonists
Grooming Development behaviors Development behaviors
Rat Cat Mouse
Decreased Delayed Decreased
6 5 18
3S 53 25 1131
392 204 596
VP VP VP
Barrel rolling Flank marking Flank marking
Rat Hamster Hamster
Increased Increased Increased
7 14 17
213 1049 1359
290 244 5
VP
Flank marking
Hamster
Increased
16
269
4
VP VP
Grooming Grooming and scratching
Rat Rat
Increased Increased
18 2
1389 1
153 350
VP VP
Hind limb movement Scratching
Rat Gerbil
Decreased Increased
9 4
1335 801
324 311
Dorsal periaqueductal gray matter admin.
Neonates Neonates ICV but not SC; in fasted, but not satiated, animals SC but not ICV Neonates Admin. during embryogenesis; neonates ICV Medial preoptic area admin. Medial preoptic-anterior hypothalamus admin.; estradiol- and progesteronesensitive response Medial preoptic-anterior hypothalamus, central gray or lateral septum-bed nucleus admin.; testosterone-sensitive effect ICV With central, but not peripheral, admin.; in nonstressful, but not stressful, conditions Intrathecal admin. With IV, but not SC, admin.; spontaneously seizing gerbil
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2233
Table 7 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
VP
Scratching
Rat
Increased
13
17
547
VP VP and OT Z-L-glutamyl-L-leucine
Vocalization Grooming Jumping
Sheep Rat Mouse
Decreased Increased Delayed
9 15 4
1221 229 417
417 52 295
Z-L-glycyl-L-proline
Jumping
Mouse
Delayed
4
417
295
Z-L-leucyl-L-glycine
Jumping
Mouse
No change
4
417
295
Z-L-prolyl-L-leucine
Jumping
Mouse
No change
4
417
295
Z-L-prolyl-D-leucine
Jumping
Mouse
Delayed
Intrathecal admin.; potentiated by naltrexone ICV Prenatal exposure Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping
4
417
295
2234
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 8 Stereotyped and other behaviors sorted by behavior Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
AT II
Amphetamine-induced stereotypy Amphetamine-induced stereotypy
Rat
Increased
ICV
9
475
58
Rat
Decreased
2
9
285
Rat
Increased
9
475
58
CCK CCK analog Boc-CCK-4
Amphetamine-induced stereotypy Anxiety behavior Anxiety behavior
Effect antagonized by 2-bromo-ergocryptine and naloxone ICV
Rat Rat
Increased No change
16 19
1313 27
263 247
CCK frag. CCK-8
Anxiety behavior
Rat
No change
19
27
247
Galanin MCH
Anxiety behavior Anxiety behavior
Mouse Rat
No change Decreased
16 17
1283 171
556 207
MSH (␣)
Anxiety behavior
Rat
Increased
17
171
207
NPY Pancreatic polypeptide ACTH frag. (1–24) ACTH frag. (4–10) NPY
Anxiety Anxiety Anxiety Anxiety Anxiety
Rat Rat Chicken Chicken Mouse
Decreased No change Increased Increased Decreased at low dose, increased at high dose
909 1445 915 915 359
428 14 414 414 387
AT II
Apomorphine-induced stereotypy Apomorphine-induced stereotypy Apomorphine-induced stereotypy Barrel rolling Barrel rolling
Rat
Increased
ICV; anxiolytic effect via Y1 receptors and anxiogenic via Y2 receptors ICV
14 20 11 11 19
9
475
58
Rat
Increased
ICV
9
475
58
Rat
Decreased
8
39
26
Rat Mouse
Increased Increased
8 15
1089 607
201 579
OT Somatostatin Somatostatin frag. (3–8), (9–14), or (7–10) VP BBS Gastrin-releasing peptide Kassinin Neuromedin B Ranatensin SP Substance K SP
Barrel rolling Barrel rolling Barrel rolling
Rat Rat Rat
Increased Increased No change
Rats treated chronically with haloperidol ICV or IV With admin. to ventrolateral thalamus but not other brain areas ICV ICV ICV
6 10 10
747 1153 1153
492 569 569
Barrel rolling Bite/scratch behavior Bite/scratch behavior Bite/scratch behavior Bite/scratch behavior Bite/scratch behavior Bite/scratch behavior Bite/scratch behavior Biting
Rat Mouse Mouse Mouse Mouse Mouse Mouse Mouse Mouse
Increased Increased Increased Increased Increased Increased Increased Increased Increased
7 7 7 7 7 7 7 7 20
213 835 835 835 835 835 835 835 301
290 47 47 47 47 47 47 47 469
Tachykinin agonist GR73632 CRF
Biting
Mouse
Increased
20
301
469
Boxing behavior
Rat
Decreased
6
891
107
SP
Boxing behavior
Rat
Increased
6
891
107
Endorphin (␥)
Sarlasin
Saralasin Cyclo (Leu-Gly) Dynorphin frag. (1–13) NPY antibody
behavior behavior behaviors behaviors behaviors
Central nucleus of the amygdala admin. Central nucleus of the amygdala admin. ICV Admin. to medial preoptic area Admin. to medial preoptic area ICV ICV
ICV Intraspinal admin. Intraspinal admin. Intraspinal admin. Intraspinal admin. Intraspinal admin. Intraspinal admin. Intraspinal admin. Spinal subarachnoid admin. Spinal subarachnoid admin. Medial frontal cortex admin.; carbacholinduced boxing behavior Medial frontal cortex admin.; carbacholinduced boxing behavior
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2235
Table 8 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
SP antagonist [D-Pro , D-Phe7, D-Trp9]-SP SP antagonist [D-Pro2, D-Trp7,9]-SP SP antisera
Boxing behavior
Rat
Decreased
9
1S 117
521
Boxing behavior
Rat
Decreased
9
1S 117
521
Boxing behavior
Rat
No change
9
1S 117
521
CCK-8
Catalepsy
Rat
Blocked
6
237
253
Ceruletide
Catalepsy
Rat
Decreased
10
779
250
Ceruletide analogs
Catalepsy
Mouse
Increased
3
701
610
Dermorphin
Catalepsy
Rat
Increased
6
3S 165
59
Dermorphin
Catalepsy
Rat
Decreased
6
3S 165
59
Endorphin ()
Catalepsy
Rat
Increased
6
237
253
MIF-1
Catalepsy
Mouse
Prevented tolerance
8
1051
380
MIF-1
Catalepsy
Rat
Decreased
2
105
85
MIF-1 analog parepide
Catalepsy
Rat
Increased
5
883
381
MIF-1 analog YPLG
Catalepsy
Mouse
No change
8
1051
380
NPY NPY frag. (1–30) SP CCK-4 CCK-8 CCK-8, desulfated Dynorphin frag. (1–6) Dynorphin frag. (1–7) Dynorphin frag. (1–8)
Catalepsy Catalepsy Chafing movements Circling Circling Circling Circling Circling Circling
Rat Rat Fish Rat Rat Rat Rat Rat Rat
Increased No effect Increased No change Increased No change Increased Increased Increased
Carbachol-induced boxing behavior Carbachol-induced boxing behavior Carbachol-induced boxing behavior ICV; endorphininduced catalepsy; effect blocked by frontal cortex lesion Haloperidol-induced catalepsy Hexobarbital-induced sleep ICV; effects increased by caudate lesion ICV; in dermorphintolerant, but not morphine-tolerant, rats ICV; effects enhanced by frontal cortex ablation Haloperidol-induced catalepsy Haloperidol-induced catalepsy Haloperidol-induced catalepsy Haloperidol-induced catalepsy ICV ICV
15 15 6 7 7 7 8 8 8
799 799 139 809 809 809 837 837 837
54 54 86 367 367 367 186 186 186
Dynorphin frag. (2–17) Enkephalin (Leu) MIF-1
Circling Circling Circling
Rat Rat Rat
Increased Increased No change
8 8 2
837 837 189
186 186 148
NPY antibody
Circling
Mouse
Increased
15
607
579
Ceruletide analogs
Convulsions
Mouse
Decreased
3
701
610
CCK-4
Defecation
Rat
No change
6
91
275
CCK-8
Defecation
Rat
No change
6
91
275
MIF-1
Defecation
Rat
Decreased
2
1S 143
120
MSH
Defecation
Rat
Increased
1
147
119
2
ICV ICV ICV Substantia nigra admin. Substantia nigra admin. Substantia nigra admin.; not antagonized by naloxone Substantia nigra admin. Substantia nigra admin. ICV; dopamine- or 5HT-stimulated circling With admin. to ventromedial thalamus, but not other brain areas Harman-induced convulsions Nucleus accumbens admin. Nucleus accumbens admin. Novelty-induced defecation Passive-avoidanceinduced defecation
(continued on next page)
2236
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 8 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
MSH
Defecation
Rat
Increased
2
1S 143
120
Secretin MIF-1 Vasotocin VIP antagonists
Defecation Depressive behaviors Development behaviors Development behaviors
Rat Human Cat Mouse
Increased Decreased Delayed Decreased
Novelty-induced defecation ICV
4 4 5 18
739 297 25 1131
83 559 204 596
ACTH ACTH frag. (4–10) Casomorphin frag. (1–5) Endorphin (␣)
Distress Distress Distress Distress
vocalization vocalization vocalization vocalization
Chicken Chicken Chicken Chicken
Increased Increased Decreased Decreased
11 11 5 5
915 915 829 823
414 414 415 573
Endorphin ()
Distress vocalization
Chicken
Decreased
5
823
573
Endorphin (␥)
Distress vocalization
Chicken
Decreased
5
823
573
VP
Flank marking
Hamster
Increased
14
1049
244
VP
Flank marking
Hamster
Increased
17
1359
5
VP
Flank marking
Hamster
Increased
16
269
4
CRF antagonist ␣-helical CRF Galanin
Freezing behavior
Rat
Decreased
15
1303
237
Freezing behavior
Rat
No change
15
1303
237
Ceruletide analogs
Gnawing
Mouse
Increased
2
2S 65
609
Secretin NPY NPY frag. (1–30) ACTH
Gnawing Grasping Grasping Grooming
Rat Rat Rat Rat
No change Increased No effect Increased
4 15 15 7
739 799 799 597
83 54 54 476
ACTH ACTH
Grooming Grooming
Rat Rat
Increased Increased
4 5
907 713
202 546
ACTH ACTH
Grooming Grooming
Rat Rat
Increased Increased
6 7
369 651
562 160
ACTH
Grooming
Rat
Increased
16
1263
434
ACTH
Grooming
Rat and mouse
Increased
8
841
159
ACTH frag. (1–24)
Grooming
Rat
Increased
4
833
6
ACTH frag. (1–4)
Grooming
Rat
Enhanced
7
1
134
ACTH/MSH frag. analogs, C-frag. elongated or cyclized
Grooming
Rat
Increased
5
1197
232
Neonates Admin. during embryogenesis; neonates
ICV; chicks With ICV, but not IP, admin. With ICV, but not IP, admin.; chicks With ICV, but not IP, admin. Medial preoptic area admin. Medial preopticanterior hypothalamus admin.; estradiol- and progesterone-sensitive response Medial preopticanterior hypothalamus, central gray or lateral septum-bed nucleus admin.; testosteronesensitive effect Foot-shock-induced behavior Foot-shock-induced behavior Methylphen-induced gnawing ICV ICV ICV ICV; blocked by naloxone ICV Pre-optic, anterior hypothalamic admin. ICV ICV; area of action localized to anteriorventral third ventricle ICV; not antagonized by nitric oxide synthase inhibitor ICV; greater effect in rats than mice ICV; effect antagonized by naloxone In MSH-induced grooming; no effect alone More potent than unmodified frag.
(continued on next page)
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
2237
Table 8 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
Amylin BBS BBS
Grooming Grooming Grooming
Rat Rat Rat
No change Increased Increased
17 2 8
589 2S 179 237
91 198 303
BBS BBS
Grooming Grooming
Rat Rat
Increased Increased
12 13
761 1215
182 99
BBS
Grooming
Rat
Increased
4
693
355
BBS
Grooming
Rat
Increased
7
557
352
BBS
Grooming
Rat
Increased
9
1S 233
260
BBS
Grooming
Rat
Increased
9
1S 245
260
BBS
Grooming
Rat
Increased
4
907
202
BBS
Grooming
Rat
Increased
6
369
562
BBS
Grooming
Rat
Increased
6
1179
564
BBS
Grooming
Rat
Increased
6
1179
563
CGRP
Grooming
Rat
Increased
17
1183
294
CGRP CRF CRF CRF
Grooming Grooming Grooming Grooming
Rat Rat Rat Rat
Decreased Increased Increased Increased
5 13 6 20
861 1149 3S 53 509
297 146 392 571
CRF
Grooming
Rat and mouse
Increased
8
841
159
Dynorphin analog desTyr1-dynorphin (1–13) Dynorphin frag. (1–13)
Grooming
Rat
Increased
4
833
6
Grooming
Rat
Increased
4
833
6
Dynorphin frag. (1–13)
Grooming
Rat and mouse
Increased
1
341
578
Galanin
Grooming
Rat
No change
11
1001
304
MSH MSH (␣)
Grooming Grooming
Rat Rat
Increased Increased
2 2
333 101
401 402
MSH (␣) MSH (␣)
Grooming Grooming
Rat Rat
Increased Increased
ICV ICV With admin. to multiple brain areas ICV ICV; developed over 8 days ICV; effect antagonized by neuroleptics ICV; antagonized by morphine With admin. to nucleus tractus solitarius but not nucleus accumbens With admin. to nucleus tractus solitarius but not nucleus accumbens With ICV, but not IV, admin.; no antagonism by haloperidol, morphine, naloxone, or NT. No effect of hypophysectomy or adrenalectomy ICV; different doses induce different types of grooming Effect enhanced by haloperidol, antagonized by naloxone and NT ICV; antagonized by haloperidol Antagonized by nitric oxide synthase inhibitor ICV ICV ICV but not SC ICV; not antagonized by melacortin-4 receptor antagonist ICV; greater effect in rats than mice; not antagonized by dexamethasone ICV; effect antagonized by naloxone ICV; effect antagonized by naloxone ICV; not reversible by naloxone Paraventicular nucleus admin. ICV Not antagonized by ␥ MSH ICV ICV; antagonized by diazepam or baclofen
4 16
721 821
286 110
(continued on next page)
2238
R.N. McLay et al. / Peptides 22 (2001) 2181–2255
Table 8 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
MSH (␣)
Grooming
Rat
Increased
18
393
470
MSH (␣)
Grooming
Rat
Increased
7
1
134
MSH (␣)
Grooming
Rat
Increased
ICV; antagonized by melanin-concentrating hormone Enhanced by ACTH frag. (1–4) Ventral tegmental area admin.; effect enhanced by bicuculline, blocked by atropine
12
203
127
MSH (␥) MSH analog [Nle4, D-Phe7]-␣ MSH MSH analogs NP-E-I
Grooming Grooming
Rat Rat
No change Increased
2 4
101 721
402 286
Grooming Grooming
Rat Rat
Increased Increased
6 18
1185 393
505 470
NT OT OT
Grooming Grooming Grooming
Rat Rat Rat
No change Increased Increased
4 12 9
493 113 1s 223
511 405 522
OT Sauvagine
Grooming Grooming
Rat Rat
Increased Increased
18 6
1389 3S 53
153 392
Sauvagine Secretin SP
Grooming Grooming Grooming
Rat Rat Mouse
Decreased No change Increased
6 4 6
3S 53 739 363
392 83 217
SP SP
Grooming Grooming
Rat Rat
Increased Increased
1 7
103 557
262 352
SP frag. (1–7) SP frag. analog pyroglutamyl-SP (7–11) Tachykinin antagonist [Sar9,Met(O2)11]SP TK PG-KII TRH
Grooming Grooming
Mouse Mouse
Decreased Increased
4 4
763 763
219 219
Grooming
Rat
Increased
18
1349
87
Grooming Grooming
Rat Rat
Increased Increased
18 9
825 283
437 565
TRH analog RGH 2202 TRH analog RX 77386 Urotensin I
Grooming Grooming Grooming
Rat Rat Rat
Increased Increased Increased
12 11 6
1309 897 3S 53
152 258 392
Urotensin I VP VP and OT BBS
Grooming Grooming Grooming Grooming and scratching
Rat Rat Rat Rat
Decreased Increased Increased Increased
6 18 15 10
3S 53 1389 229 529
392 153 52 259
ICV
ICV; antagonized by MSH PVN and IV admin. ICV Ventral tegmental area admin.; effect not altered by sex, sexsteroids, or dopamine antagonists ICV ICV but not SC; in fasted, but not satiated, animals SC but not ICV ICV ICV; different effectiveness in different strains and ages; enhanced by naloxone but not phenoxybenzamine or haloperidol ICV ICV; not antagonized by morphine
Nucleus basalis magnocellularis admin. ICV ICV; effect antagonized by haloperidol, naloxone and NT Neonates ICV but not SC; in fasted, but not satiated, animals SC but not ICV ICV Prenatal exposure Neonates
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2239
Table 8 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
OT
Grooming and scratching
Rat
Increased
2
1
350
VP
Grooming and scratching
Rat
Increased
2
1
350
MIF-1
Head turning
Rat
No change
2
189
148
MIF-1 VP Dynorphin frag. (1–13) CRF
Mouse Rat Rat Monkey
No change Decreased Increased Increased
2 9 4 4
189 1335 631 217
148 324 168 268
Casomorphin analog BCH 325 L-pyroglutamyl-Lleucine Z-L-glutamyl-L-leucine
Head twitch Hind limb movement Hind limb paralysis Huddling and lying down Jerking
With central, but not peripheral, admin.; in non-stressful, but not stressful, conditions With central, but not peripheral, admin.; in non-stressful, but not stressful, conditions Striatally-evoked head turning 5HT-stimulated twitch Intrathecal admin. Intrathecal admin. ICV
Rat
Decreased
16
635
447
Jumping
Mouse
Delayed
4
417
295
Jumping
Mouse
Delayed
4
417
295
Z-L-glycyl-L-proline
Jumping
Mouse
Delayed
4
417
295
Z-L-leucyl-L-glycine
Jumping
Mouse
No change
4
417
295
Z-L-prolyl-L-leucine
Jumping
Mouse
No change
4
417
295
Z-L-prolyl-D-leucine
Jumping
Mouse
Delayed
Bromocriptine-induced jerks Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping Morphine-withdrawalinduced jumping
4
417
295
BBS BBS Gastrin-releasing peptide
Licking Licking Licking
Rat Rat Rat
Decreased Decreased Decreased
17 16 17
107 903 107
175 523 175
Gastrin-releasing peptide Neuromedin B-10
Licking Licking
Rat Rat
Decreased Decreased
16 17
903 107
523 175
SP
Licking
Mouse
Increased
20
301
469
Tachykinin agonist GR73632 TRH
Licking
Mouse
Increased
20
301
469
Licking
Rat
Increased
2
1S 99
187
Morphine-induced behaviors Myoclonus (serotonergic behavior) Obsessive compulsive behavior Opiate withdrawal behavior Oral act. (serotonergic behavior)
Mouse
No change
17
1291
97
Mouse
No change
10
5
439
Human
Decreased
13
1083
139
Rat
No change
15
1303
237
Rat
No change
14
1159
206
Mouse Rat Rat
Increased Increased Increased
2 4 4
2S 65 277 277
609 395 395
Adrenal Peptide E (Frog) ACTH OT Galanin MIF-1
Ceruletide analogs BBS Calcitonin
Ptosis Respiration Respiration
Admin. simultaneously with neuromedin B-10 Admin. simultaneously with gastrin-releasing peptide Spinal subarachnoid admin. Spinal subarachnoid admin. ICV; punished licking in water-deprived, ethanol-treated rats ICV 5-hydroxytryptophaninduced In a single patient
6-hydroxydopaminelesioned rats treated with desipramine ICV ICV
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Table 8 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
CCK Enkephalin analog DADLE Enkephalin analog DADLE Enkephalin analog DAGO Enkephalin analog FK33–824 NT NT SP TRH
Respiration Respiration
Cat Rat
Increased tidal volume Increased
9 3
809 1031
246 225
Respiration
Rat
Increased
Ventral medulla admin. Nucleus tractus solitarius admin. Third ventricle admin.
3
1023
236
Respiration
Rat
Nucleus tractus solitarius admin.
3
1031
225
Respiration
Mouse
Increased at low dose decreased at high dose Decreased rate
9
777
378
Respiration Respiration Respiration Respiration
Monkey Rat Rat Rat
Decreased No change No change Increased
5 4 4 4
125 277 277 277
366 395 395 395
BBS
Righting
Rat
Impaired
2
1S 99
187
Endorphin ()
Righting
Rat
Impaired
2
1S 99
187
NT
Righting
Rat
Impaired
2
1S 99
187
TRH
Righting
Rat
Improved
2
1S 99
187
SP
Rigidity
Rat
Reversed
1
103
262
SP
Risk assessment behavior
Rat
Increased
20
1437
126
SP frag. (7–11)
Risk assessment behavior
Rat
Increased
20
1437
126
ACTH/MSH 4–10 analog ORG 2766
Rotation
Rat
Decreased
14
1317
10
CCK-8
Rotation
Rat
Increased
18
1161
396
Cyclo (Leu-Gly)
Rotation
Rat
No change
8
39
26
MIF-1 analog 3(R)-[(2(S)pyrrolidinylcarbonyl) amino]-2-oxo-1pyrrolidineacetamide NT SP
Rotation
Rat
Increased
ICV ICV ICV ICV; decreased tidal volume ICV; alcohol-induced righting impairment ICV; alcohol-induced righting impairment ICV; alcohol-induced righting impairment ICV; alcohol-induced righting impairment ICV; in haloperidol treated Dorsal periaqueductal gray matter admin. Dorsal periaqueductal gray matter admin. Nigrostriatal lesions and amphetamineinduced turning Animals pre-treated with peptide prior to nigrostriatal lesions; dopamine-depletioninduced turning Apomorphine-induced rotation in rats with unilateral nigrostriatal lesions Apomorphine- or LDOPA-induced rotation in 6-hydroxydopaminelesioned rats
18
1209
363
Rotation Rotation
Rat Rat
Increased Normalized
6 18
1057 1161
389 396
CCK-8 Gastrin (1–17) BBS
Rumination Rumination Scratching
Sheep Sheep Mouse
No change Increased Increased
5 5 9
81 81 909
238 238 567
BBS
Scratching
Rat
Increased
6
1179
563
Substantia nigra admin. Animals pre-treated with peptide prior to nigrostriatal lesions; dopamine-depletioninduced turning ICV Intrathecal admin.; not antagonized by DPro2,D-Trp7,9-SP and phenoxybenzamine ICV; antagonized by naloxone
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2241
Table 8 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
Somatostatin
Scratching
Mouse
Increased
9
909
567
SP SP
Scratching Scratching
Mouse Mouse
Increased Increased
4 6
517 363
249 217
SP
Scratching
Mouse
Increased
9
909
567
SP
Scratching
Mouse
Increased
20
301
469
Tachykinin agonist GR73632 VP
Scratching
Mouse
Increased
20
301
469
Scratching
Gerbil
Increased
4
801
311
VP
Scratching
Rat
Increased
13
17
547
SP Dynorphin frag. (1–13) Enkephalin (Met) Enkephalin analog DAMA TRH TRH analog RX 77386 BBS MIF-1 analog Cyclo(Leu-Gly) Endorphin () Melatonin OT CCK
Scratching and licking Shaking Shaking Shaking
Rat Rat Fish Rat
Increased Increased Increased Increased
Intrathecal admin.; not antagonized by DPro2,D-Trp7,9-SP and phenoxybenzamine Intrathecal admin. ICV; different effectiveness in different strains and ages Intrathecal admin.; antagonized by DPro2,D-Trp7,9-SP and phenoxybenzamine Spinal subarachnoid admin. Spinal subarachnoid admin. With IV, but not SC, admin.; spontaneously seizing gerbil Intrathecal admin.; potentiated by naltrexone Intrathecal admin. ICV or IV
3 8 6 4
49 1089 139 239
485 201 86 155
Shaking Shaking Sighing Sniffing
Rat Rat Rat Rat
Increased Increased Increased Increased
4 11 4 5
239 897 277 7
155 258 395 310
Startle Startle response Startle response Stretching
Rat Rat Rat Mouse
Decreased Decreased Increased Decreased
2 2 6 10
1S 137 1S 155 567 31
234 188 284 9
OT TRH Endorphin ()
Tail movement Tail movement Trembling
Rat Rat Rat
Increased Increased Decreased
6 6 2
1191 1191 1S 99
538 538 187
Enkephalin MSH
Vocalization Vocalization
Sheep Chicken
Decreased Decreased
9 11
1221 647
417 413
VP ACTH AT BBS Bradykinin NT OT SP MSH antagonist [D-Trp7, Ala8, DPhe10)␣-MSH (6–11)amide Calcitonin
Vocalization Vomiting Vomiting Vomiting Vomiting Vomiting Vomiting Vomiting Walking pattern
Sheep Dog Dog Dog Dog Dog Dog Dog Rat
Decreased No change No change No change No change Increased Increased No change Increased recovery
9 6 6 6 6 6 6 6 16
1221 1S 173 1S 173 1S 173 1S 173 1S 173 1S 173 1S 173 319
417 597 597 597 597 597 597 597 433
Writhing
Mouse
Increased analgesia
6
3S 277
200
ICV ICV Neonates ICV Apomorphine-induced Sniffing
ICV; abdominal irritant-induced stretching Intrathecal admin. Intrathecal admin. ICV; alcohol-induced tremors ICV Separation-induced vocalization in chicks ICV ICV ICV ICV ICV ICV ICV ICV Admin. after sciatic nerve crush
ICV, IP, and IV, but no effect SC; Achinduced writhing
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Table 8 (continued) Peptide
Behavior
Species
Effect
Notes
Vol
Page
Ref
ACTH
Yawning
Rat
Increased
16
1263
434
Casomorphin analog BCH 325
Yawning
Rat
Normalized
16
635
447
Casomorphin analogs
Yawning
Rat
Decreased
13
69
464
NT
Yawning
Rat
Decreased
12
755
398
NT analog[D-Trp11]NT
Yawning
Rat
Decreased
12
755
398
OT
Yawning
Rat
Increased
ICV; antagonized by nitric oxide synthase inhibitor Bromocriptine-evoked yawning that was inhibited by reserpine ICV; apomorphineinduced yawning ICV; apomorphineinduced yawning ICV; apomorphineinduced yawning ICV
10
559
11
2.7. Stereotyped and other behaviors No categorization system is likely to capture all of the types of behaviors that peptides can affect. Because readers might be interested in a particular behavior listed in this catch-all category, the table is presented twice, once organized by peptide (Table 7) and once by behavior (Table 8).
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R.N. McLay et al. / Peptides 22 (2001) 2181–2255 [117] Danguir J. Food intake in rats is increased by intracerebroventricular infusion of the somatostatin analogue SMS 201–995 and is decreased by somatostatin antiserum. Peptides 1988;9:211–3. [118] Datta PC, Hoehler FK, Sandman CA. Effects of melatonin on startle reflex in rat. Peptides 1981;2 Suppl 1:155– 60. [119] Datta PC, King MG. Effects of ␣-MSH and melatonin on passive avoidance and on PA-induced defecation and plasma 11-OHCS in hypophysectomized rats. Peptides 1980;1:147–53. [120] Datta PC, King MG. ␣-MSH, MIF-I, and melatonin: effects on novelty-induced defecation, plasma 11-OHCS, and central catecholamines in rats. Peptides 1981;2 Suppl 1:143–54. [121] Datta PC, Sandman CA, Hoehler FK. Attenuation of morphine analgesia by ␣-MSH, MIF-I, melatonin and naloxone in the rat. Peptides 1982;3:433–7. [122] Davis JL, Pico RM. Arginine vasotocin delays extinction of a conditioned avoidance behavior in neonatal chicks. Peptides 1984; 5:1221–3. [123] Davis JL, Pico RM, Cherkin A. L-prolyl-L-arginyl-glycinamide induces memory enhancement in chicks. Peptides 1983;4:401– 4. [124] Davis JM, Lowy MT, Yim GK, Lamb DR, Malven PV. Relationship between plasma concentrations of immunoreactive -endorphin and food intake in rats. Peptides 1983;4:79 – 83. [125] Davis TP, Gillespie TJ, Porreca F. Peptide fragments derived from the -chain of hemoglobin (hemorphins) are centrally active in vivo. Peptides 1989;10:747–51. [126] De Araujo JE, Silva RC, Huston JP, Brandao ML. Anxiogenic effects of substance P and its 7–11 C terminal, but not the 1–7 N terminal, injected into the dorsal periaqueductal gray. Peptides 1999; 20:1437– 43. [127] de Barioglio SR, Lezcano N, Celis ME. Alpha MSH-induced excessive grooming behavior involves a GABAergic mechanism. Peptides 1991;12:203–5. [128] de Beaurepaire R, Suaudeau C. Anorectic effect of calcitonin, neurotensin and bombesin infused in the area of the rostral part of the nucleus of the tractus solitarius in the rat. Peptides 1988;9:729 –33. [129] De Caro G, Massi M. Water intake modifications induced by tachykinins, bombesins and opioid peptides. Peptides 1985;6(Suppl 3): 181–5. [130] De Caro G, Massi M, Micossi LG, Perfumi M. Angiotensin II antagonists versus drinking induced by bombesin or eledoisin in pigeons. Peptides 1982;3:631– 6. [131] De Caro G, Massi M, Micossi LG, Perfumi M. Drinking and feeding inhibition by ICV pulse injection or infusion of bombesin, ranatensin and litorin to rats. Peptides 1984;5:607–13. [132] de Castiglione R. Structural requirements of ceruletide-like peptides for activity on gut and brain. Peptides 1981;2 Suppl 2:61–3. [133] De Ceballos ML, Lopez AE, Harto JR, Bravo A, Gomez-Monterrey I, Gonzalez-Muniz R, Garcia-Lopez MT, Del Rio J. Prolonged antinociceptive activity of pseudodipeptide analogues of LysTrp(Nps) and Trp(Nps)-Lys. Peptides 1992;13:63–7. [134] De Graan PN, Spruijt BM, Eberle AN, Girard J, Gispen WH. ACTH1– 4 potentiates ␣-MSH-induced melanophore dispersion, and excessive grooming. Peptides 1986;7:1– 4. [135] de Pedro N, Cespedes MV, Delgado MJ, Alonso-Bedate M. Muopioid receptor is involved in -endorphin-induced feeding in goldfish. Peptides 1996;17:421– 4. [136] de Pedro N, Pinillos ML, Valenciano AI, Alonso-Bedate M, Delgado MJ. Inhibitory effect of serotonin on feeding behavior in goldfish: involvement of CRF. Peptides 1998;19:505–11. [137] Della-Fera MA, Baile CA. CCK-octapeptide injected in CSF decreases meal size, and daily food intake in sheep. Peptides 1980;1: 51– 4. [138] Della-Fera MA, Baile CA, Beinfeld MC. Cerebral ventricular transport and uptake: importance for CCK-mediated satiety. Peptides 1982;3:963– 8. [139] den Boer JA, Westenberg HG. Oxytocin in obsessive compulsive disorder. Peptides 1992;13:1083–5.
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[140] Denbow DM. Centrally and peripherally administered bombesin decreases food intake in turkeys. Peptides 1989;10:275–9. [141] Denbow DM, Duke GE, Chaplin SB. Food intake, gastric secretion, and motility as affected by avian pancreatic polypeptide administered centrally in chickens. Peptides 1988;9:449 –54. [142] Denbow DM, Myers RD. Eating, drinking, and temperature responses to intracerebroventricular cholecystokinin in the chick. Peptides 1982;3:739 – 43. [143] Deupree D, Hsiao S. Cholecystokinin octapeptide, proglumide, and passive avoidance in rats. Peptides 1987;8:25– 8. [144] Deutch AY, Maggio JE, Bannon MJ, Kalivas PW, Tam SY, Goldstein M, Roth RH. Substance K, and substance P differentially modulate mesolimbic, and mesocortical systems. Peptides 6 Suppl 1985;2:113–22. [145] Deviche P, Schepers G. Intracerebroventricular injection of ostrich -endorphin to satiated pigeons induces hyperphagia but not hyperdipsia. Peptides 1984;5:691– 4. [146] Diamant M, Croiset G, De Wied D. The effect of corticotropinreleasing factor (CRF) on autonomic and behavioral responses during shock-prod burying test in rats. Peptides 1992;13:1149 –58. [147] Dickinson SL, Slater P. Opiate receptor antagonism by L-prolyl-Lleucyl-glycinamide, MIF-I. Peptides 1980;1:293–9. [148] Dickinson SL, Slater P. Effect of L-prolyl-L-leucyl-glycinamide (MIF-I) on some neurotransmitter-receptor interactions. Peptides 1981;2:189 –95. [149] Dickson PR, Feifel D, Vaccarino FJ. Blockade of endogenous GRF at dark onset selectively suppresses protein intake. Peptides 1995; 16:7–9. [150] Dickson PR, Vaccarino FJ. GRF-induced feeding: evidence for protein selectivity, and opiate involvement. Peptides 1994;15:1343– 52. [151] Dluzen DE, Muraoka S, Engelmann M, Landgraf R. The effects of infusion of arginine vasopressin, oxytocin, or their antagonists into the olfactory bulb upon social recognition responses in male rats. Peptides 1998;19:999 –1005. [152] Drago F, Grassi M, Valerio C, Coppi G, Lauria N, Nicotra GC, Raffaele R. Behavioral changes induced by the thyrotropin-releasing hormone analogue, RGH 2202. Peptides 1991;12:1309 –13. [153] Drago F, Stanciu M, Salehi S, Scapagnini U. The block of central vasopressin V1 but not V2 receptors suppresses grooming behavior and hypothermia induced by intracerebroventricular vasopressin in male rats. Peptides 1997;18:1389 –92. [154] Drucker-Colin R, Bernal-Pedraza J, Fernandez-Cancino F, Oksenberg A. Is vasoactive intestinal polypeptide (VIP) a sleep factor? Peptides 1984;5:837– 40. [155] Drust EG, Crawford IL. Comparison of the effects of TRH and D-Ala2-metenkephalinamide on hippocampal electrical activity and behavior in the unanesthetized rat. Peptides 1983;4:239 – 43. [156] Dryden S, Pickavance L, Henderson L, Williams G. Hyperphagia induced by hypoglycemia in rats is independent of leptin and hypothalamic neuropeptide Y (NPY). Peptides 1998;19:1549 –55. [157] Dudley CA, Vale W, Rivier J, Moss RL. The effect of LHRH antagonist analogs and an antibody to LHRH on mating behavior in female rats. Peptides 1981;2:393– 6. [158] Dulawa SC, Vanderweele DA. Cholecystokinin and estradiol synergistically potentiate satiety in rats. Peptides 1994;15:913– 8. [159] Dunn AJ, Berridge CW, Lai YI, Yachabach TL. CRF-induced excessive grooming behavior in rats, and mice. Peptides 1987;8:841– 4. [160] Dunn AJ, Hurd RW. ACTH acts via an anterior ventral third ventricular site to elicit grooming behavior. Peptides 1986;7:651–7. [161] Eaves M, Thatcher-Britton K, Rivier J, Vale W, Koob GF. Effects of corticotropin releasing factor on locomotor activity in hypophysectomized rats. Peptides 1985;6:923– 6. [162] Eckel LA, Geary N. Endogenous cholecystokinin’s satiating action increases during estrus in female rats. Peptides 1999;20:451– 6.
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