The Rat Brain. In Stereotaxic Coordinates

262 Pages·1986·50.7 MB·English

by  George Paxinos (Auth.)

The Rat Brain
in Stereotaxic Coordinates
Second Edition
George Paxinos
School of Psychology
University of New South Wales
Charles Watson
Department of Health
Perth, Western Australia
ACADEMIC PRESS
(Harcourt Brace Jovanovich, Publishers)
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ACADEMIC PRESS AUSTRALIA
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Copyright © 1986 by
ACADEMIC PRESS AUSTRALIA
All rights reserved. No part of this pubhcation may be
reproduced or transmitted in any form or by any means,
electronic or mechanical, including photocopy, recording,
or any information storage and retrieval system, without
permission in writing from the publisher.
Printed in Australia
National Library of Australia Cataloguing-in-Publication Data
Paxinos, George, date .
The rat brain in stereotaxic coordinates.
2nd ed.
Bibliography.
Includes index.
ISBN 0 12 547621 3.
1. Rats — Anatomy — Atlases. 2. Brain — Atlases.
I. Watson, Charles, date . II. Title.
599.32*33
Library of Congress Catalog Card Number: 86-71486
To William R. Mehler in appreciation of his willingness to
share his formidable knowledge of neuroanatomy.
Preface: Second Edition that our publishers will attend promptly to any written request significance of the differential distribution of neuroactive
to reproduce the figures in this atlas. However, we would like to compounds.
take this opportunity to stress that electrode placements and We thank Natalie Chabin, Anita Soekarno and Robert
The publication of the second edition allows us the opportunity lesions can be represented accurately on atlas drawings only Hutchison for their skill and outstanding dedication in
to expand The Rat Brain in Stereotaxic Coordinates in order to after careful inspection of sections taken from the brain of the improving our ink drawings and ensuring that errors in labeling
provide a more comprehensive guide for those interested in the experimental subject. were kept to a minimum. We are grateful to Louise Kahabka for
rat brain. The plates and diagrams of the first edition were not at excellent typing of the manuscript and to Elly Paxinos for
close enough intervals to effectively represent all major brain editorial assistance. We also thank Sharlane Velasco, Paul
structures. In addition, the subdivisions of many brain areas Acknowledgements Nolan, Vera Thomson, Brian Jeffreys, Maureen Lodsman, and
were not mapped in sufficient detail. Anna Watson for technical assistance.
The second edition differs from the first in that: We also wish to thank the staff of Academic Press Australia
1. It has nearly twice as many plates and accompanying Second Edition for the exceptional quality of reproduction of our work.
Original work reported here was supported by Australian NH
diagrams.
& MRC grants to George Paxinos.
2. There is an increase in magnification of (a) the coronal We wish to express our deep appreciation to scientists who
Finally, and most importantly, we wish to thank Elly and
plates caudal to the interaural line, (b) the sagittal, and (c) generously assisted us in the construction of the second edition.
Prue, our wives, for tolerating our extended stays in the
the horizontal plates. Our greatest debt is to William R. Mehler who shared with us
laboratory for the last seven months.
3. It has nearly twice as many structures labeled. the rich experiences of his exploration of the thalamus and the
4. It includes cortical parcellations according to Zilles brainstem.
(1985). We are also deeply indebted to Clifford B. Saper for critical
5. Its delineations and nomenclature have benefited from the advice on the hypothalamus, parabrachial nuclei, solitary First Edition
scrutiny of colleagues expert in different brain regions. nucleus, and insular cortex.
6. A more expanded **basis of delineations" is given. One of the most significant features of the present edition is We are grateful to a number of scientists who took hours, even
the inclusion of the comprehensive cortical parcellation of Karl days, from their busy schedules to identify structures on the
We have gratefully accepted the suggestions of our colleagues Zilles. Zilles' (1985) book is a milestone in cortical Plates and correct boundaries we had drawn. Our interaction
on delineations and nomenclature. We have been tremendously neuroanatomy and we owe a special thanks to him for allowing with them and their challenge to our preconceptions were the
assisted by the authors who contributed to The Rat Nervous us to use his plan. most rewarding experiences of this project. With pleasure we
System (Paxinos, 1985b). This book, which complements the We express our deep appreciation to the invaluable advice thank Paula Wilson (for help with the pons and medulla). Bill
atlas, enabled us to undertake a more sophisticated analysis in given to us by Jose de Olmos (amygdala and bed nucleus of the Mehler (thalamus and midbrain), Jim Fallon (amygdala, ventral
the second edition of this atlas. stria terminalis); Kirsten Osen (inferior coUiculus, superior tegmental area, raphe nuclei and locus coeruleus), Richard Faull
While the usual copyright restrictions pertain for the olive, and cochlear nuclei); Jim Fallon (brainstem and basal (thalamus and substantia nigra), Jan Meyer (amygdala and
reproduction of figures of this atlas. Academic Press Australia forebrain); Ruth Bleier (hypothalamus); Laszlo Zaborszky and construction of abbreviations), Alan Brichta (spinal cord),
will respond promptly to written requests to reproduce specific Lennart Heimer (accumbens, ventral pallidum, and medial Istvan Tork (ventral tegmental area and raphe nuclei), John
sets of figures. We have one request to make of users of this forebrain bundle); Stanley W^eigand (preoptic area); Barbara Haight (neocortex, thalamus and construction of
atlas: In the interest of facilitating communication, please Jones (reticular formation); Joseph LeDoux (medial geniculate); abbreviations), Konrad Talbot (delineations in AChE-stained
consider the suitability of our system of nomenclature and David Tracey (nucleus Ζ and trigeminal nuclei); Henk sections), Lennart Heimer, Robert Switzer and Joanna Hill
abbreviations for your work. This system is a distillation of Groenewegen (interpeduncular and mediodorsal nuclei); Alvin (ventral pallidum and olfactory peduncle), Larry Swanson
views from a number of neuroscientists. Beitz (central gray); Terry Takahashi (laterodorsal and (hypothalamus and ventral tegmental area), Shirley Bayer
lateroposterior thalamic nuclei); Yasuhiro Torigoe (interstitial (olfactory bulb and hippocampal formation), Steve McHanwell
nucleus of the medial longitudinal fasciculus and reticular (lumbar and sacral spinal cord). Maxwell Cowan (hippocampus
Preface: First Edition
tegmental nucleus); Bill Armstrong (paraventricular and mammillary bodies), John Morrison (neocortex), George
hypothalamic nucleus); Masaya Tohyama (habenula and dorsal Martin (medulla), Paul MacLean (forebrain), Sandra Loughlin
tegmental nuclei); Suzanne Haber (globus pallidus); Richard (neocortex), Esmail Meisami (olfactory bulb), Dennis Steindler
The mapping of the central nervous system is an art pioneered Faull (thalamus); George Martin (reticular formation); Kent and Irena Grofova (substantia nigra), Steven Hunt (Rexed's
by the great neuroanatomists of the nineteenth century. The Morest (auditory system); Jeffery Winer and Kate Games layers), Derek van der Kooy and Leonard Koda (medulla),
challenge of recent times has been to place such maps in an (subbrachial nucleus); David Amaral (nomenclature); Brent Irving Diamond (thalamus), John Johnson (thalamus), Bill
accurate stereotaxic reference system and to revise the Vogt (cortex); Mary-Clare Hoist (inferior olive). Armstrong (hypothalamus), Sharleen Sakai (thalamus), Fritz
delineation of cell groups and fiber tracts in the light of the latest Our concepts on the organization of the brainstem have Guldner (suprachiasmatic nucleus), Eva Bystrzycka (KoUiker-
findings. benefited from a long standing interaction with our colleague Fuse nucleus), Wilfred Schober (for making available to us his
In an effort to construct an atlas of horizontal sections of the Istvan Tork. In addition, we wish to thank him for allowing us personal copy of the Wunscher, Schober and Werner atlas), Phil
rat forebrain (Paxinos et αι., 1980), we noticed that brain to view his collection of 5-hydroxytryptamine and substance Ρ Marks, Paul Herron and Mick O'Brien (for checking stereotaxic
sections demonstrated enhanced staining contrast and suffered stained brainstem sections. accuracy), and Piers Emson (for making available brain sections
less distortion if they were cut from an unfixed brain. We Because of the opportunity to review the excellent Timm's from DFP-treated rats).
thought that it might be possible to obtain better delineation of stained preparations of Gorm Danscher (Danscher, 1984; We are indebted to Professor Syd Lovibond and Professor
nuclei and achieve greater stereotaxic accuracy by using Danscher and Norgaard, 1985) we improved our boundaries in a Fred Rost for making available facilities of the Schools of
fresh-frozen brains, especially if the brains were frozen while number of areas especially the ventral pallidum and substantia Psychology and Anatomy at the University of New South
still in the skull. We were also aware of the need for an atlas innominata. We also wish to thank him for allowing us to Wales, and to Professor Nicole Le Douarin for providing
based on the convenient flat-skull position, suitable for adult include unpublished observations (Paxinos and Danscher) on laboratory space at the Institut d'Embryologie at Nogent-sur-
male rats, and which took into account the anatomical research the existence of the terete and striohypothalamic nuclei. Marne.
of the last two decades. Such an atlas, we felt, should also Larry Butcher made available to G. Paxinos AChE-stained We express our gratitude to Alan Brichta for constructing the
represent sections from all regions of the central nervous system. sections following DFP pretreatment in relation to an earlier skull diagram.
The Rat Brain in Stereotaxic Coordinates is intended for use collaborative study (Paxinos and Butcher, 1985). This material, We are particularly grateful to Gordana Strumfin and Ann
by researchers and graduate students in the neurosciences. In stained in accordance with the excellent procedures developed by Topple for their excellent technical assistance in photography,
addition, senior undergraduates should find the atlas a useful Butcher, was critical for the identification of nuclei presented in drawing of the figures and construction of figure labels, and to
adjunct to readings and lectures in brain anatomy and function. that study and has enhanced our appreciation of the use of Lorraine Brooks for typing the manuscript. Our sincere thanks
The photographs are of sufficient magnification to permit chemoarchitecture as a guide to the organization of regions. also to the following for assisting us in important ways in the
researchers to make their own judgments on the boundaries of We thank Charles Ribak for allowing us to view sections production of the histological material, plates and drawings:
brain structures. We would be pleased to receive any comments stained for glutamic acid decarboxylase. After seeing clearly the Elly Paxinos, Sharon McDonald, Julia Watson, Geoff
on the delineations we have made so that future editions may subdivisions of the zona incerta in his material we proceeded to Schneider, Janelle Hopwood, Kevin Maynard and Maree
benefit from the experience of our colleagues. outline them in our plates. Garrett.
Although reproduction of any part of this book is subject to We thank Claudio Cuello with whom we have enjoyed a
the usual restrictions of copyright, we can assure researchers lengthy collaboration and shared an appreciation of the
Vll
Preface: Second Edition that our publishers will attend promptly to any written request significance of the differential distribution of neuroactive
to reproduce the figures in this atlas. However, we would like to compounds.
take this opportunity to stress that electrode placements and We thank Natalie Chabin, Anita Soekarno and Robert
The publication of the second edition allows us the opportunity lesions can be represented accurately on atlas drawings only Hutchison for their skill and outstanding dedication in
to expand The Rat Brain in Stereotaxic Coordinates in order to after careful inspection of sections taken from the brain of the improving our ink drawings and ensuring that errors in labeling
provide a more comprehensive guide for those interested in the experimental subject. were kept to a minimum. We are grateful to Louise Kahabka for
rat brain. The plates and diagrams of the first edition were not at excellent typing of the manuscript and to Elly Paxinos for
close enough intervals to effectively represent all major brain editorial assistance. We also thank Sharlane Velasco, Paul
structures. In addition, the subdivisions of many brain areas Acknowledgements Nolan, Vera Thomson, Brian Jeffreys, Maureen Lodsman, and
were not mapped in sufficient detail. Anna Watson for technical assistance.
The second edition differs from the first in that: We also wish to thank the staff of Academic Press Australia
1. It has nearly twice as many plates and accompanying Second Edition for the exceptional quality of reproduction of our work.
Original work reported here was supported by Australian NH
diagrams.
& MRC grants to George Paxinos.
2. There is an increase in magnification of (a) the coronal We wish to express our deep appreciation to scientists who
Finally, and most importantly, we wish to thank Elly and
plates caudal to the interaural line, (b) the sagittal, and (c) generously assisted us in the construction of the second edition.
Prue, our wives, for tolerating our extended stays in the
the horizontal plates. Our greatest debt is to William R. Mehler who shared with us
laboratory for the last seven months.
3. It has nearly twice as many structures labeled. the rich experiences of his exploration of the thalamus and the
4. It includes cortical parcellations according to Zilles brainstem.
(1985). We are also deeply indebted to Clifford B. Saper for critical
5. Its delineations and nomenclature have benefited from the advice on the hypothalamus, parabrachial nuclei, solitary First Edition
scrutiny of colleagues expert in different brain regions. nucleus, and insular cortex.
6. A more expanded **basis of delineations" is given. One of the most significant features of the present edition is We are grateful to a number of scientists who took hours, even
the inclusion of the comprehensive cortical parcellation of Karl days, from their busy schedules to identify structures on the
We have gratefully accepted the suggestions of our colleagues Zilles. Zilles' (1985) book is a milestone in cortical Plates and correct boundaries we had drawn. Our interaction
on delineations and nomenclature. We have been tremendously neuroanatomy and we owe a special thanks to him for allowing with them and their challenge to our preconceptions were the
assisted by the authors who contributed to The Rat Nervous us to use his plan. most rewarding experiences of this project. With pleasure we
System (Paxinos, 1985b). This book, which complements the We express our deep appreciation to the invaluable advice thank Paula Wilson (for help with the pons and medulla). Bill
atlas, enabled us to undertake a more sophisticated analysis in given to us by Jose de Olmos (amygdala and bed nucleus of the Mehler (thalamus and midbrain), Jim Fallon (amygdala, ventral
the second edition of this atlas. stria terminalis); Kirsten Osen (inferior coUiculus, superior tegmental area, raphe nuclei and locus coeruleus), Richard Faull
While the usual copyright restrictions pertain for the olive, and cochlear nuclei); Jim Fallon (brainstem and basal (thalamus and substantia nigra), Jan Meyer (amygdala and
reproduction of figures of this atlas. Academic Press Australia forebrain); Ruth Bleier (hypothalamus); Laszlo Zaborszky and construction of abbreviations), Alan Brichta (spinal cord),
will respond promptly to written requests to reproduce specific Lennart Heimer (accumbens, ventral pallidum, and medial Istvan Tork (ventral tegmental area and raphe nuclei), John
sets of figures. We have one request to make of users of this forebrain bundle); Stanley W^eigand (preoptic area); Barbara Haight (neocortex, thalamus and construction of
atlas: In the interest of facilitating communication, please Jones (reticular formation); Joseph LeDoux (medial geniculate); abbreviations), Konrad Talbot (delineations in AChE-stained
consider the suitability of our system of nomenclature and David Tracey (nucleus Ζ and trigeminal nuclei); Henk sections), Lennart Heimer, Robert Switzer and Joanna Hill
abbreviations for your work. This system is a distillation of Groenewegen (interpeduncular and mediodorsal nuclei); Alvin (ventral pallidum and olfactory peduncle), Larry Swanson
views from a number of neuroscientists. Beitz (central gray); Terry Takahashi (laterodorsal and (hypothalamus and ventral tegmental area), Shirley Bayer
lateroposterior thalamic nuclei); Yasuhiro Torigoe (interstitial (olfactory bulb and hippocampal formation), Steve McHanwell
nucleus of the medial longitudinal fasciculus and reticular (lumbar and sacral spinal cord). Maxwell Cowan (hippocampus
Preface: First Edition
tegmental nucleus); Bill Armstrong (paraventricular and mammillary bodies), John Morrison (neocortex), George
hypothalamic nucleus); Masaya Tohyama (habenula and dorsal Martin (medulla), Paul MacLean (forebrain), Sandra Loughlin
tegmental nuclei); Suzanne Haber (globus pallidus); Richard (neocortex), Esmail Meisami (olfactory bulb), Dennis Steindler
The mapping of the central nervous system is an art pioneered Faull (thalamus); George Martin (reticular formation); Kent and Irena Grofova (substantia nigra), Steven Hunt (Rexed's
by the great neuroanatomists of the nineteenth century. The Morest (auditory system); Jeffery Winer and Kate Games layers), Derek van der Kooy and Leonard Koda (medulla),
challenge of recent times has been to place such maps in an (subbrachial nucleus); David Amaral (nomenclature); Brent Irving Diamond (thalamus), John Johnson (thalamus), Bill
accurate stereotaxic reference system and to revise the Vogt (cortex); Mary-Clare Hoist (inferior olive). Armstrong (hypothalamus), Sharleen Sakai (thalamus), Fritz
delineation of cell groups and fiber tracts in the light of the latest Our concepts on the organization of the brainstem have Guldner (suprachiasmatic nucleus), Eva Bystrzycka (KoUiker-
findings. benefited from a long standing interaction with our colleague Fuse nucleus), Wilfred Schober (for making available to us his
In an effort to construct an atlas of horizontal sections of the Istvan Tork. In addition, we wish to thank him for allowing us personal copy of the Wunscher, Schober and Werner atlas), Phil
rat forebrain (Paxinos et αι., 1980), we noticed that brain to view his collection of 5-hydroxytryptamine and substance Ρ Marks, Paul Herron and Mick O'Brien (for checking stereotaxic
sections demonstrated enhanced staining contrast and suffered stained brainstem sections. accuracy), and Piers Emson (for making available brain sections
less distortion if they were cut from an unfixed brain. We Because of the opportunity to review the excellent Timm's from DFP-treated rats).
thought that it might be possible to obtain better delineation of stained preparations of Gorm Danscher (Danscher, 1984; We are indebted to Professor Syd Lovibond and Professor
nuclei and achieve greater stereotaxic accuracy by using Danscher and Norgaard, 1985) we improved our boundaries in a Fred Rost for making available facilities of the Schools of
fresh-frozen brains, especially if the brains were frozen while number of areas especially the ventral pallidum and substantia Psychology and Anatomy at the University of New South
still in the skull. We were also aware of the need for an atlas innominata. We also wish to thank him for allowing us to Wales, and to Professor Nicole Le Douarin for providing
based on the convenient flat-skull position, suitable for adult include unpublished observations (Paxinos and Danscher) on laboratory space at the Institut d'Embryologie at Nogent-sur-
male rats, and which took into account the anatomical research the existence of the terete and striohypothalamic nuclei. Marne.
of the last two decades. Such an atlas, we felt, should also Larry Butcher made available to G. Paxinos AChE-stained We express our gratitude to Alan Brichta for constructing the
represent sections from all regions of the central nervous system. sections following DFP pretreatment in relation to an earlier skull diagram.
The Rat Brain in Stereotaxic Coordinates is intended for use collaborative study (Paxinos and Butcher, 1985). This material, We are particularly grateful to Gordana Strumfin and Ann
by researchers and graduate students in the neurosciences. In stained in accordance with the excellent procedures developed by Topple for their excellent technical assistance in photography,
addition, senior undergraduates should find the atlas a useful Butcher, was critical for the identification of nuclei presented in drawing of the figures and construction of figure labels, and to
adjunct to readings and lectures in brain anatomy and function. that study and has enhanced our appreciation of the use of Lorraine Brooks for typing the manuscript. Our sincere thanks
The photographs are of sufficient magnification to permit chemoarchitecture as a guide to the organization of regions. also to the following for assisting us in important ways in the
researchers to make their own judgments on the boundaries of We thank Charles Ribak for allowing us to view sections production of the histological material, plates and drawings:
brain structures. We would be pleased to receive any comments stained for glutamic acid decarboxylase. After seeing clearly the Elly Paxinos, Sharon McDonald, Julia Watson, Geoff
on the delineations we have made so that future editions may subdivisions of the zona incerta in his material we proceeded to Schneider, Janelle Hopwood, Kevin Maynard and Maree
benefit from the experience of our colleagues. outline them in our plates. Garrett.
Although reproduction of any part of this book is subject to We thank Claudio Cuello with whom we have enjoyed a
the usual restrictions of copyright, we can assure researchers lengthy collaboration and shared an appreciation of the
Vll
Preface: Second Edition that our publishers will attend promptly to any written request significance of the differential distribution of neuroactive
to reproduce the figures in this atlas. However, we would like to compounds.
take this opportunity to stress that electrode placements and We thank Natalie Chabin, Anita Soekarno and Robert
The publication of the second edition allows us the opportunity lesions can be represented accurately on atlas drawings only Hutchison for their skill and outstanding dedication in
to expand The Rat Brain in Stereotaxic Coordinates in order to after careful inspection of sections taken from the brain of the improving our ink drawings and ensuring that errors in labeling
provide a more comprehensive guide for those interested in the experimental subject. were kept to a minimum. We are grateful to Louise Kahabka for
rat brain. The plates and diagrams of the first edition were not at excellent typing of the manuscript and to Elly Paxinos for
close enough intervals to effectively represent all major brain editorial assistance. We also thank Sharlane Velasco, Paul
structures. In addition, the subdivisions of many brain areas Acknowledgements Nolan, Vera Thomson, Brian Jeffreys, Maureen Lodsman, and
were not mapped in sufficient detail. Anna Watson for technical assistance.
The second edition differs from the first in that: We also wish to thank the staff of Academic Press Australia
1. It has nearly twice as many plates and accompanying Second Edition for the exceptional quality of reproduction of our work.
Original work reported here was supported by Australian NH
diagrams.
& MRC grants to George Paxinos.
2. There is an increase in magnification of (a) the coronal We wish to express our deep appreciation to scientists who
Finally, and most importantly, we wish to thank Elly and
plates caudal to the interaural line, (b) the sagittal, and (c) generously assisted us in the construction of the second edition.
Prue, our wives, for tolerating our extended stays in the
the horizontal plates. Our greatest debt is to William R. Mehler who shared with us
laboratory for the last seven months.
3. It has nearly twice as many structures labeled. the rich experiences of his exploration of the thalamus and the
4. It includes cortical parcellations according to Zilles brainstem.
(1985). We are also deeply indebted to Clifford B. Saper for critical
5. Its delineations and nomenclature have benefited from the advice on the hypothalamus, parabrachial nuclei, solitary First Edition
scrutiny of colleagues expert in different brain regions. nucleus, and insular cortex.
6. A more expanded **basis of delineations" is given. One of the most significant features of the present edition is We are grateful to a number of scientists who took hours, even
the inclusion of the comprehensive cortical parcellation of Karl days, from their busy schedules to identify structures on the
We have gratefully accepted the suggestions of our colleagues Zilles. Zilles' (1985) book is a milestone in cortical Plates and correct boundaries we had drawn. Our interaction
on delineations and nomenclature. We have been tremendously neuroanatomy and we owe a special thanks to him for allowing with them and their challenge to our preconceptions were the
assisted by the authors who contributed to The Rat Nervous us to use his plan. most rewarding experiences of this project. With pleasure we
System (Paxinos, 1985b). This book, which complements the We express our deep appreciation to the invaluable advice thank Paula Wilson (for help with the pons and medulla). Bill
atlas, enabled us to undertake a more sophisticated analysis in given to us by Jose de Olmos (amygdala and bed nucleus of the Mehler (thalamus and midbrain), Jim Fallon (amygdala, ventral
the second edition of this atlas. stria terminalis); Kirsten Osen (inferior coUiculus, superior tegmental area, raphe nuclei and locus coeruleus), Richard Faull
While the usual copyright restrictions pertain for the olive, and cochlear nuclei); Jim Fallon (brainstem and basal (thalamus and substantia nigra), Jan Meyer (amygdala and
reproduction of figures of this atlas. Academic Press Australia forebrain); Ruth Bleier (hypothalamus); Laszlo Zaborszky and construction of abbreviations), Alan Brichta (spinal cord),
will respond promptly to written requests to reproduce specific Lennart Heimer (accumbens, ventral pallidum, and medial Istvan Tork (ventral tegmental area and raphe nuclei), John
sets of figures. We have one request to make of users of this forebrain bundle); Stanley W^eigand (preoptic area); Barbara Haight (neocortex, thalamus and construction of
atlas: In the interest of facilitating communication, please Jones (reticular formation); Joseph LeDoux (medial geniculate); abbreviations), Konrad Talbot (delineations in AChE-stained
consider the suitability of our system of nomenclature and David Tracey (nucleus Ζ and trigeminal nuclei); Henk sections), Lennart Heimer, Robert Switzer and Joanna Hill
abbreviations for your work. This system is a distillation of Groenewegen (interpeduncular and mediodorsal nuclei); Alvin (ventral pallidum and olfactory peduncle), Larry Swanson
views from a number of neuroscientists. Beitz (central gray); Terry Takahashi (laterodorsal and (hypothalamus and ventral tegmental area), Shirley Bayer
lateroposterior thalamic nuclei); Yasuhiro Torigoe (interstitial (olfactory bulb and hippocampal formation), Steve McHanwell
nucleus of the medial longitudinal fasciculus and reticular (lumbar and sacral spinal cord). Maxwell Cowan (hippocampus
Preface: First Edition
tegmental nucleus); Bill Armstrong (paraventricular and mammillary bodies), John Morrison (neocortex), George
hypothalamic nucleus); Masaya Tohyama (habenula and dorsal Martin (medulla), Paul MacLean (forebrain), Sandra Loughlin
tegmental nuclei); Suzanne Haber (globus pallidus); Richard (neocortex), Esmail Meisami (olfactory bulb), Dennis Steindler
The mapping of the central nervous system is an art pioneered Faull (thalamus); George Martin (reticular formation); Kent and Irena Grofova (substantia nigra), Steven Hunt (Rexed's
by the great neuroanatomists of the nineteenth century. The Morest (auditory system); Jeffery Winer and Kate Games layers), Derek van der Kooy and Leonard Koda (medulla),
challenge of recent times has been to place such maps in an (subbrachial nucleus); David Amaral (nomenclature); Brent Irving Diamond (thalamus), John Johnson (thalamus), Bill
accurate stereotaxic reference system and to revise the Vogt (cortex); Mary-Clare Hoist (inferior olive). Armstrong (hypothalamus), Sharleen Sakai (thalamus), Fritz
delineation of cell groups and fiber tracts in the light of the latest Our concepts on the organization of the brainstem have Guldner (suprachiasmatic nucleus), Eva Bystrzycka (KoUiker-
findings. benefited from a long standing interaction with our colleague Fuse nucleus), Wilfred Schober (for making available to us his
In an effort to construct an atlas of horizontal sections of the Istvan Tork. In addition, we wish to thank him for allowing us personal copy of the Wunscher, Schober and Werner atlas), Phil
rat forebrain (Paxinos et αι., 1980), we noticed that brain to view his collection of 5-hydroxytryptamine and substance Ρ Marks, Paul Herron and Mick O'Brien (for checking stereotaxic
sections demonstrated enhanced staining contrast and suffered stained brainstem sections. accuracy), and Piers Emson (for making available brain sections
less distortion if they were cut from an unfixed brain. We Because of the opportunity to review the excellent Timm's from DFP-treated rats).
thought that it might be possible to obtain better delineation of stained preparations of Gorm Danscher (Danscher, 1984; We are indebted to Professor Syd Lovibond and Professor
nuclei and achieve greater stereotaxic accuracy by using Danscher and Norgaard, 1985) we improved our boundaries in a Fred Rost for making available facilities of the Schools of
fresh-frozen brains, especially if the brains were frozen while number of areas especially the ventral pallidum and substantia Psychology and Anatomy at the University of New South
still in the skull. We were also aware of the need for an atlas innominata. We also wish to thank him for allowing us to Wales, and to Professor Nicole Le Douarin for providing
based on the convenient flat-skull position, suitable for adult include unpublished observations (Paxinos and Danscher) on laboratory space at the Institut d'Embryologie at Nogent-sur-
male rats, and which took into account the anatomical research the existence of the terete and striohypothalamic nuclei. Marne.
of the last two decades. Such an atlas, we felt, should also Larry Butcher made available to G. Paxinos AChE-stained We express our gratitude to Alan Brichta for constructing the
represent sections from all regions of the central nervous system. sections following DFP pretreatment in relation to an earlier skull diagram.
The Rat Brain in Stereotaxic Coordinates is intended for use collaborative study (Paxinos and Butcher, 1985). This material, We are particularly grateful to Gordana Strumfin and Ann
by researchers and graduate students in the neurosciences. In stained in accordance with the excellent procedures developed by Topple for their excellent technical assistance in photography,
addition, senior undergraduates should find the atlas a useful Butcher, was critical for the identification of nuclei presented in drawing of the figures and construction of figure labels, and to
adjunct to readings and lectures in brain anatomy and function. that study and has enhanced our appreciation of the use of Lorraine Brooks for typing the manuscript. Our sincere thanks
The photographs are of sufficient magnification to permit chemoarchitecture as a guide to the organization of regions. also to the following for assisting us in important ways in the
researchers to make their own judgments on the boundaries of We thank Charles Ribak for allowing us to view sections production of the histological material, plates and drawings:
brain structures. We would be pleased to receive any comments stained for glutamic acid decarboxylase. After seeing clearly the Elly Paxinos, Sharon McDonald, Julia Watson, Geoff
on the delineations we have made so that future editions may subdivisions of the zona incerta in his material we proceeded to Schneider, Janelle Hopwood, Kevin Maynard and Maree
benefit from the experience of our colleagues. outline them in our plates. Garrett.
Although reproduction of any part of this book is subject to We thank Claudio Cuello with whom we have enjoyed a
the usual restrictions of copyright, we can assure researchers lengthy collaboration and shared an appreciation of the
Vll
Vlll
Introduction 2. The atlas is based on the study of 130 adult male Wistar characteristic dendritic architecture (e.g. the subnuclei of the
rats (with a weight range of 270-310 g). It is suitable for superior olive, the inferior colliculus, the mediodorsal thalamic
brains of 250-350 g male and female rats. nucleus, and the central amygdaloid nucleus). Although we often
For a long time the rat has been widely used as a subject for 3. It represents all areas of the brain and spinal cord, and referred to a set of brain sections in which fibers were stained
research in the neurosciences. This is chiefly because of its size, brain areas are shown in coronal, sagittal and horizontal with luxol fast blue, we found the sections stained for AChE of
resistance to infection and the availability of inbred strains. In planes. The brain sections shown were taken on average at much greater value for the delineation of nuclei and fiber tracts.
recent years, the use of rats has increased significantly, 0.25 mm intervals and were stained with either cresyl
particularly in the field of neurohistochemistry. With increasing violet or for the demonstration of acetylcholinesterase Surgery
public pressure against the use of medium-sized mammals (the (AChE).
monkey, cat and rabbit) in experimental biology, many 4. It is based on fresh brains frozen in the skull (using deeply Under barbiturate anaesthesia the rats were placed in a Kopf
laboratories have opted to use only small rodents. While the anaesthetized rats) in order to overcome distortion small-animal stereotaxic instrument, and the incisor bar was
mouse is ideal for studies involving development and genetic produced by fixation and to enhance staining contrast. adjusted until the heights of lambda and bregma skull points
selection (because of its short generation time), the rat, having a 5. Structures are delineated on the basis of data on were equal. This flat-skull position was achieved when the
larger brain, is more appropriate for accurate stereotaxic cytoarchitecture, chemoarchitecture, and connectivity. incisor bar was lowered 3.3 ± 0.4 mm below horizontal zero
localization of discrete brain areas. (Table 1). Because the point of intersection of the lambdoid and
In spite of the pre-eminent popularity of the rat for stereotaxic sagittal sutures is variable, we have chosen to define lambda as
procedures, there is no atlas of the rat brain based on the Why acetylkcholinesterase? the midpoint of the curve of best fit along the lambdoid suture
convenient flat-skull position that is sufficiently comprehensive (see skull diagram). This redefined reference point is
and stereotaxically accurate for adult (250-350 g) male rats. In This atlas grew out of our efforts to make an atlas of horizontal considerably more reliable than the true lambda, and it is
addition, no current atlas has photographs of sufficient sections of the rat forebrain in stereotaxic coordinates (Paxinos located 0.3 ± 0.3 mm anterior to the interaural line. We also
magnification to permit investigators to judge for themselves the et al., 1980). We chose the acetylcholinesterase (AChE) stain for defined bregma as the point of intersection of the sagittal suture
veracity of the atlas delineations. that atlas for two reasons. Firstly, AChE clearly delineates many with the curve of best fit along the coronal suture. When the two
Before beginning the construction of the first editon of the nuclei and fiber tracts whose boundaries are difficult to sides of the coronal suture met the sagittal suture at different
present atlas, we surveyed the 181 articles reporting use of distinguish in Nissl-stained or Weil-stained sections. Secondly, points, bregma usually fell midway between the two junctions.
stereotaxic surgery in rats which were published during 1980 in the use of AChE staining provides an indication of the extent of The anteroposterior position of bregma was 9.1 ± 0.3 mm
three prominent neuroscience journals (Brain Research, damage to fiber systems that contain acetylcholine or the anterior to the coronal plane passing through the interaural line.
Neuroscience Letters and Physiology and Behavior). We did this monoamines (Butcher and Marchand, 1978; Emson et al., Since the mean anteroposterior position of bregma for the three
in order to assess current preferences in stereotaxic methods and 1979). Furthermore, it has been shown that the distribution of rats from which the photographs were taken was slightly less
nomenclature. In 52^70 of the studies surveyed, the Konig and AChE in certain areas is in register with the distribution of a than 9.1 mm, 9.0 mm was used on the atlas diagrams. The top of
Klippel (1963) atlas was used; in 20Ψο, the Pellegrino and number of neuropeptides (Graybiel et al., 1981). the skull at bregma and lambda was 10.0 ± 0.2 mm dorsal to the
Cushman atlas was used (Pellegrino and Cushman 1967; We have found that the use of AChE-stained sections interaural zero plane. Table 1 presents mean body-weight and
Pellegrino et al., 1979); and in the remainder, other atlases were alongside Nissl-stained sections is invaluable for the delineation mean skull measurements for all rats as well as individual
used. In 79^o of the studies, male rats were used, and the mean of many nuclei (e.g. the superior vestibular, the olfactory measurements for the rats from which the plate photographs
weight was 276 g. In 63% of the studies, bregma was used as the tubercle, the basolateral amygdaloid, and the ventral thalamic were taken.
reference point; in 25%, the interaural line; and in 12%, nuclei). In addition, the use of this combination of stains makes To establish the stereotaxic position of structures, reference
lambda. In 85% of the reports, English or anglicized possible the subdivision of certain nuclei into subnuclei that needle tracks were made perpendicular to the horizontal,
nomenclature was used for the naming of the brain structures. correspond to regions with particular afferent patterns or coronal and sagittal planes. For brains sectioned in the coronal
Because the Konig and Klippel atlas is by far the most widely
used atlas of the rat brain, we took note of the advantages of this
atlas in the preparation of our own. We believe that the Konig
and Klippel atlas is popular with researchers because its
reference system closely approximates the flat-skull position and Table 1
brain structures are correctly identified in almost every case. Craniometric and stereotaxic data (means ± S.D.) for rats of different sex, strain and weight.
However, since the publication of Konig and Klippel's
excellent atlas, knowledge of the anatomy of brain structures Mean AP AP DV AP AP AP DV
wweeiigghhtt I-B I-L I-B I-Acb B-ac I-7n I - incisor bar
has increased substantially, and the accepted parcelling of such Subject ((gg))** (mm) (mm) (mm) (mm)** (mm)** (mm)** (mm)
regions as the thalamus, septum and amygdala has changed
significantly. In addition, some major brain areas currently 'Atlas'
studied with stereotaxic surgery (the pons, medulla and Wistar 290 9.1 ±0.3 0.3 ±0.3 10.0±0.2 11.7 0.0 -1.3 -3.3±0.4
cerebellum) are not represented in the Konig and Klippel atlas. A Coronal
plates 300 9.2 0.2 10.1
further consideration is that the Konig and Klippel atlas is based
Sagittal
on brains of 150 g female rats, whereas in 1980 more than two-
plates 270 8.9 0.0 10.0
thirds of researchers used 250-350 g male rats. This difference in
Horizontal
size and sex results in an error of over 2.0 mm in the
plates 290 9.1 0.2 10.1
anteroposterior coordinate of many structures (Paxinos et al., Female
1980), and many researchers have found it necessary to Wistar 282 9.3 ±0.2 0.5 ±0.3 10.0±0.1 11.6 0.1 -1.2 -3.2±0.5
determine empirically the coordinates of each structure of Hooded 290 9.4 ±0.4 0.3 ±0.6 9.8 ±0.2 11.9 0.0 -1.2 -3.9±0.6
interest. Finally, the Konig and Klippel atlas does not give the Sprague 299 9.0±0.2 0.7±0.2 10.1±0.1 11.7 0.1 -1.2 -3.9±0.5
position of bregma, and the drawings of brain structures do not Juvenile
match the photographs presented. Wistar 180 7.7±0.4 -0.4±0.3 9.9±0.2 10.2 -0.1 -1.6 -2.0±0.4
Mature
The present work represents our efforts to meet the challenge
Wistar 436 9.7±0.3 0.6 ±0.3 10.7 ±0.4 12.4 -0.1 -0.8 -2.7±0.3
of providing an atlas of sufficient accuracy, comprehensiveness
and convenience for the needs of present-day neuroscientists.
*S.D.s. < 20 g.
The main features of this atlas are: •*S.D.s. < 0.4 mm.
ac, anterior commissure; Acb, accumbens nucleus; AP, anterior-posterior; B, bregma; DV, dorsal-ventral; 7n, facial nerve; I, interaural line; L, lambda.
1. It is based on the flat-skull position, and bregma, lambda,
Reprinted with permission from J. Neuroscience Methods. 13 (1985) 139-143.
or the midpoint of the interaural line can be used as a
reference point.
plane, vertical needle insertions were made at 2.0 mm intervals Cresyl violet was drawn in after consideration of sections obtained from
through the brain (except for the penetrations at 0.7 mm decalcified heads. These heads were sectioned with brains in situ
anterior to the interaural line, which was chosen to avoid Slides were immersed for 5 min in each of the following: xylene, and stained with cresyl violet and luxol fast blue.
rupture of a venous sinus). Ten such needle tracks appear on xylene, 100<^o alcohol, lOO^o, 95Ψο and 70^o alcohol. They were Fiber tracts were outlined by solid lines and nuclei by dashed
coronal plates of this atlas. Three horizontal needle insertions dipped in distilled water (H2O) and stained in 0.5% cresyl violet lines. In general, the abbreviations are placed in the center of
perpendicular to the coronal plane were made from the posterior for 15-30 min. They were differentiated in H2O for 3-5 min and structures. For the coronal and horizontal drawings, the
of the brain at 1.0, 3.0 and 5.0 mm above the interaural line and then dehydrated through 70%, 95%, 100% and 100% alcohol. abbreviations for fiber tracts and fissures are almost always
approximately 1.0 mm lateral to the midline. The reference They were then put in xylene and cover-slipped. positioned on the left hemisphere of the figure, and for nuclei
tracks from the horizontal needles appear as pinholes in coronal To make 500 mL of 0.5% cresyl violet of about pH 3.9, mix they are generally positioned on the right hemisphere. The
sections. (a) 2.5 g of cresylecht violet (Chroma Gesellschaft); (b) 300 mL ventricles are filled in with solid black.
For brains sectioned in the sagittal plane, vertical needles were of H2O; (c) 30 mL of 1.0 Μ sodium acetate (13.6 g of granular
inserted in both hemispheres at 3.0 mm posterior to the soidum acetate in 92 mL of H2O); and (d) 170 mL of 1.0 Μ Reference planes
interaural line and at 1.0 and 2.0 mm lateral to the midline. A acetic acid (29 mL of glacial acetic acid added to 471 mL of
second pair was inserted 11 mm anterior to the interaural line at H2O). Mix solution for at least 7 days on a magnetic stirrer, then Two coronal and two horizontal zero planes are given, one set
1.0 and 2.0 mm lateral to the midline. Horizontal needle tracks filter. related to the interaural line and one to bregma. Lambda was
perpendicular to the coronal plane were made at 5.0 mm and 6.0 located 0.3 ± 0.3 mm anterior to the interaural Hne, and it too
mm dorsal to the interaural line and 2.0 mm lateral to the AChE histochemistry can be used as a reference point in conjunction with the
midline. Horizontal needle tracks perpendicular to the sagittal dorsoventral coordinate of bregma. The position of the
plane were made at 5.0 mm dorsal to the interaural Hne and 2.0 The method for the demonstration of AChE followed the stereotaxic reference points and planes are indicated on the skull
mm and 8.0 mm anterior to it. procedures of Koelle and Friedenwald (1949) and Lewis (1961). diagram. The stereotaxic reference grid shows 0.2 mm intervals.
For brains sectioned in the horizontal plane, two vertical Slides were incubated for 15 h in the following solution: 100 mL Stereotaxic coordinates can be readily obtained by using a
reference tracks were made by inserting a needle at 2.0 and 9.0 of stock solution to which had been added 116 mg of transparent ruler or by superimposing a transparent grid over
mm anterior to the interaural line and approximately 1.4 mm S-acetylthiocholine iodide and 3.0 mg ethopropazine (May & the drawing.
lateral to the midline. These reference tracks appear as pinholes Baker). The slides were rinsed with tap water and developed for
in the horizontal sections. Five horizontal needle tracks 10 min in 1% sodium sulphide (1.0 g in 100 mL of H2O) at pH Coronal drawings
perpendicular to the coronal plane were made by inserting 7.5. They were then rinsed with water and immersed in 4%
needles at 1.0, 3.0, 5.0, 7.0 and 8.0 mm above the interaural paraformaldehyde in phosphate buffer for 8 h, and then allowed The large number at the bottom left of each drawing shows the
lines. to dry. Subsequently, they were dehydrated for 5 min in 100% anteroposterior distance of the corresponding plate from the
alcohol, then immersed in xylene and cover-slipped with vertical plane passing through the interaural line. The large
Histology Permount. number at bottom right shows the anteroposterior distance of
The stock solution was a 50 mM sodium acetate buffer at pH the plate from bregma. The numbers on the left margin show the
After surgery all rats were decapitated, and their brains were 5.0 which was made 4.0 mM with respect to copper sulphate and dorsoventral distance from the horizontal plane passing through
frozen on dry ice or CO2 within 3 min of decapitation. Brains 16mM with respect to glycine. (This was done by adding 6.8 g of the interaural line. The numbers on the right margin show the
that were sectioned coronally were blocked at 3.0 mm anterior to sodium acetate, 1.0 g of copper sulphate crystals and 1.2 g of dorsoventral distance from the horizontal plane passing through
the interaural line prior to freezing. Brains removed from the glycine to 1.0 L of H2O and lowering the pH to 5.0 with HCl.) bregma and lambda on the surface of the skull. The numbers on
skull for sectioning on the horizontal or sagittal planes initially It was noticed that fresh, unfixed tissue showed a substantially the top and bottom margins show the distance of structures
presented a problem in that they assumed the shape of the stage stronger reaction for both stains than tissue fixed with formalin, from the midline.
on which they were positioned. To avoid this distortion, the paraformaldehyde, glutaraldehyde or alcohol. (A detailed
brains were frozen in the skull, and the skull bones were then protocol of the staining procedures may be obtained from the Sagittal drawings
prized off the frozen brains. authors.)
Spinal cord segments were obtained from 9 rats and were The large number at the bottom left of each drawing shows the
fresh frozen in propane cooled with liquid nitrogen. Photography distance of the corresponding plate from the midline. The
Representative sections from the major regions of the spinal numbers on the left margin show the dorsoventral distance from
cord are included in the atlas. Photographs of stained brain sections were taken with a Nikon the horizontal plane passing through the interaural line. The
Frozen brains were sectioned on an American Optical Cryocut Multiplot macrophotographic apparatus on 4" χ 5" Kodak Plus numbers on the right margin show the dorsoventral distance
at 40 Mm. Sections were obtained parallel to the stereotaxic X film. High contrast (grade 4) Ilfospeed paper was used for the from the horizontal plane passing through bregma and lambda
planes by adjusting the angle of cutting until the needle tracks Nissl sections, whereas lower contrast (grade 2) paper was used on the surface of the skull. The numbers on the bottom margin
encountered were parallel (as far as was possible) to the plane of for the AChE sections. For sections in which the AChE activity show the anteroposterior distance from the coronal plane
section. Sections were taken directly from the cryotome knife on in the cerebellum and the cerebral cortex was too weak to be passing through the interaural line. The numbers on the top
uncoated slides with the help of an anti-roll device. Three clearly seen, the printing time was increased and the non-cortical margin show the anteroposterior distance from the coronal
sections at successive 0.5 mm intervals were stained alternately areas were shaded. Even after such artificial enhancement, these plane passing through bregma.
with cresyl violet or for the demonstration of AChE. One of areas still display the weakest AChE reactivity in the
these sections at each interval was presented as an atlas plate in photographs in which they appear. Horizontal drawings
the first edition of the atlas. In the second edition we have The photographs were reproduced at Macarthur Press,
interpolated an additional plate between existing plates. For the Sydney, on a Hell Computer scanner equipped with a laser light, The large number at the bottom left of each drawing shows the
coronal set the interpolated plates commence after Plate 11 and enabling an accurate halftone reproduction of 150 line screen dorsoventral distance of the corresponding plate from the
are accompanied by a labeled line drawing. For the horizontal whilst reducing the need to deep etch the unnecessary horizontal plane passing through the interaural line. The large
set, interpolated plates and drawings occur throughout the background. number at top right shows the dorsoventral distance of the plate
range. For the sagittal set, the interpolated sections (without from the horizontal plane passing through bregma and lambda
accompanying diagrams) are placed below the photographs of Drawings on the surface of the skull. The numbers on the bottom margin
the sections that are diagrammed on the facing pages. An show the anteroposterior distance from the coronal plane
attempt was made to select sections close to the midpoint of the It was thought that the drawings would be more informative if passing through the interaural line. The numbers on the top
interval of the plates in the first edition of the atlas; however they were not stylized, and for this reason artistic licence was margin show the anteroposterior distance from the coronal
such sections often were not available. Additionally, the Nissl rarely taken. There was no mirror-image drawing and, with the plane passing through bregma. The numbers on the left and
substance and AChE stained sections no longer alternate. exception of small adjustments to distorted midlines and cortex, right margins show the distance of structures from the midline.
Staining was carried out on the same day as cutting, according to the drawings depic^ the asymmetries present in the sections. Values ventral to the interaural horizontal zero or posterior to
the following procedures. When part of an atlas section was missing or severly distorted, it either the interaural line or bregma are preceded by a minus sign.
IX
An example rats (10.0 mm). In the mature rats, the interaural line to bregma used, greater accuracy can be achieved if bregma is used as the
vertical distance was 10.7 mm. reference point for work with rostral structures and the
If it is desired to insert an electrode into the basolateral In female rats, as well as in hooded, juvenile (180 g), mature interaural line for work with caudal structures. A further
amygdaloid nucleus, Fig. 29 reveals the coordinates with (436 g) and *atlas' (290 g) Wistar rats, bregma was found to be improvement in accuracy can be obtained by taking into account
reference to the interaural line to be 6.2 mm anterior to the above the most forward crossing fibers of the anterior the actual location of the accumbens nucleus (anterior part of
interaural line, 1.5 mm dorsal to it, and 5.0 mm lateral to the commissure. This is the point at which the posterior limbs of the Acb shown on Fig. 9) and the genu of the facial nerve (Fig. 59).
midline. The coordinates of the same structure obtained with anterior commissure appear. These data confirm the In agreement with Slotnick and Brown (1980), we noticed that
reference to bregma are 2.8 mm posterior to bregma, 8.5 mm observation of Whishaw et al. (1977) that bregma is more stable coordinates of structures (in rats within the size range for which
ventral to it, and 5.0mm lateral to the midUne. than the interaural line for positioning of electrodes in brain the atlas was constructed) were closer to target if the coordinates
structures close to, or anterior to, bregma. However, data from given by the interaural and bregma reference systems were
Stereotaxic accuracy insertion of needles aimed at the level where the facial nerve averaged.
leaves the facial genu (Fig. 59) show that the interaural reference In this atlas, minor discrepancies exist between the
Consideration of the actual coordinates of some structures point is more stable than bregma for localization of such coordinates obtained for a single structure from the three planes
reveals that the variability of values obtained from the three posterior structures. Therefore, if juvenile or mature rats are represented. These discrepancies result from (a) individual
planes is well within the range of values expected from the use of
three different rats. For example, the coordinates given by the
three planes for the bed nucleus of the anterior commissure and
the trochlear nucleus are similar (see Table 2). However, some
structures located far from the stereotaxic reference points may
have coordinates in the three planes differing by as much as 0.5
mm. The positioning of structures on the diagrams was not
adjusted to make the three brains conform, because such
adjustments would have removed the correspondence between
photograph and diagram.
Table 2
Coordinates of the bed nucleus of the anterior commissure and the trochlear
nucleus obtained from the three planes
Bed nucleus of the ant comm Trochlear nucleus
Plane A-P D-V Lat A-P D-V Lat
Coronal 8.2 3.4 0.9 1.7 3.4 0.4
Sagittal 8.0 3.4 0.9 1.8 3.5 0.4
Horizontal 8.1 3.4 0.9 1.8 3.4 0.4
Medium-sized (270-310 g) male rats were used in the
construction of this atlas. As a result the structures are futher
apart from each other than in the Konig and Klippel atlas, which
is based on small (150 g) female rats. Due to the difference in
size and sex, up to 2 mm differences in the location of structures
will be found if the two atlases are compared.
Atlases are frequently used for work with animals of different
sex, strain, and weight than those for which the atlases were
intended. We have estimated the error that will occur if this atlas 10.0mm
is used with female Wistar rats, male hooded (Long Evans) rats,
male Sprague Dawley rats of *atlas' weight (approximately 290
g), as well as juvenile Wistar (180 g), and mature Wistar (436 g)
rats. It is evident from Table 1 (reproduced from Paxinos et al.,
1985) that no substantial stereotaxic error will occur when rats
of different sex and strain are chosen provided that rats are of
similar weight to those on which the atlas is based (290 g). For
example, for rats of different sex and strain but of similar weight
the anteroposterior distance between the interaural line and
bregma is between 9.0 and 9.4 mm. Similarly, the dorsoventral
distance between the interaural line and the surface of the skull
at bregma and lambda is very stable (9.8-10.1 mm). By contrast,
craniometric data for juvenile (180 g) and mature (436 g) Wistar Skull diagram
rats differ substantially from those of other groups. The Dorsal and lateral views of the skull of a 290 g Wistar rat. The positions of
anteroposterior distance between the interaural line and bregma bregma, lambda and the plane of the interaural line are shown above the lateral
is 7.7 mm in the juvenile and 9.7mm in the mature rats (9.0 mm view. The distance between the horizontal plane passing through bregma and
in 'atlas' rats). Lambda is 0.4 mm posterior to the interaural line lambda and the horizontal plane passing through the interaural line is shown on
the right of the lateral view. The distance between the incisor bar and the
in the juvenile rats and 0.6 mm anterior to this line in the mature
horizontal plane passing through the interaural line is shown on the left of the
rats (0.3 mm anterior in the *atlas' rats). Unexpectedly, the
lateral view. Lambda (midpoint of the curve of best fit along the lambdoid
dorsoventral distance between the interaural line and bregma for
suture) is 0.3 mm anterior to the coronal plane passing through the interaural
juvenile rats (9.9 mm) was almost the same as that of the * atlas' line.
differences between the rats used; (b) deviation or drifts of Compound names of nuclei have a capital letter for each part the ventral pallidum (Heimer and Wilson, 1975; Heimer et al.,
needles during insertion (especially needles penetrating (e.g. MD = mediodorsal thalamic nucleus). If a word occurs in mi; Switzer et al., 1982).
horizontally for 30 mm into the brain); (c) distortion caused by the names of a number of structures, it is usually given the same
Substriatal area (SStr): Dorsal to the anterior amygdaloid area
freezing; (d) sectioning on a plane not perfectly parallel or abbreviation (e.g. reticular thalamic nucleus = Rt;
and ventral to the fundus striati lies a region which is less cellular
normal to the inserted needles; (e) distortion caused in mounting reticulotegmental nucleus of the pons = RtTg). Exceptions to
and less reactive for AChE than the fundus striati (Figs 22-24,
the sections on the shdes, and (0 constraints imposed by the this rule are made for well-established abbreviations such as
84, 93). This region is shown to be reactive for Substance Ρ (Fig.
need to obtain sections at intervals of exactly 0.5 mm. In the VTA. Abbreviations of brain regions are omitted where the
3E of Haber and Nauta, 1983) and this reactivity is continuous
coronal plates, an additional error resulted from the fact that identity of the region in question is clear from its position (e.g.
with that of the globus pallidus. Given that it lies ventral to the
Plates 1-3, 40-42 and 67-76 were taken from rat brains different Arc = arcuate hypothalamic nucleus, not ArcH). Arabic
fundus striati, we have called it the substriatal area (SStr).
from the one seen in the remaining coronal plates. This numerals are used instead of Roman numerals in identifying (a)
supplementation was necessary because of distortion of the cranial nerves and nuclei (as in the Berman, 1968, atlas), (b) Accumbens nucleus (Acb): The accumbens nucleus has been
olfactory bulbs and caudal medulla in the brain from which cortical layers, (c) cerebellar folia, and (d) layers of the spinal subdivided into an outer shell (AcbSh) and an inner core (AcbC)
most of the plates were taken and because of the loss of one cord. While the spoken meaning is the same, the detection (Figs 10-15, 78-82, 93-101). The outer shell is continuous both
section at the level at which the brain was blocked. threshold is lower, ambiguity is reduced, and they are easier to with the ventrolateral pocket of the striatum (fundus striati)
position in small spaces available on diagrams. and, through the cell bridges, with the cell dense (pyramidal)
Nomenclature layer of the olfactory tubercle. The accumbens core surrounds
The basis of delineation of structures the anterior limb of the anterior commissure and is less reactive
After our review of the literature revealed that English for AChE but more reactive in Timm's preparations than the
nomenclature is currently used in about 85% of neuroscience Nuclear and fiber-tract boundaries were determined after accumbens shell (Zaborszky et al., 1985).
articles, we decided to use English rather than Latin terms. consideration of authoritative studies published on the
Substantia innominata (SI): Following the separation of the
Retention of rarely used Latin terms and the construction of cytoarchitecture, connectivity, histochemistry and development
ventral pallidum from the substantia innominata, the remainder
latinized terms for newly named nuclei would, we believe, have of the central nervous system. Inconsistencies in demarcation
(posterior part) was called the sublenticular part of the
been an unprofitable exercise. It would also have delayed the found in the literature were usually resolved in favour of the
substantia innominata (Heimer et al., 1985; de Olmos et al.,
development of a standardized list of terms and abbreviations to delineation supported by the AChE distribution in our material.
1985). We have noticed, however, that a strip of substantia
which atlases should contribute. However, some Latin terms In many cases, assistance with the construction of boundaries
innominata persists rostrally bounded by the ventral pallidum,
that are strongly entrenched, such as *stria terminalis', have was provided by a number of colleagues who have special
the lateral preoptic area, the nucleus of the horizontal limb of
been retained and not translated into artificial anglicized forms. knowledge of the anatomy of particular brain regions (see
the diagonal band, and the magnocellular preoptic nucleus (Figs
There is a critical need for a stable neuroanatomical Acknowledgements). We also benefited from our examination
18-20, 80, 91, 92). This region displays a modest activity in
nomenclature to accurately and intelligibly convey information of the delineation of structures in a number of rat, mouse, cat
Timm's stained preparations—as does the substantia
between neuroscientists. At the present time, many terms are and human brain atlases (Berman, 1968; Berman and Jones,
innominata. We were alerted to the continuity of this 'pan­
used for the same structure, and, in some cases, the same term 1982; Meessen and Olszewski, 1949; Mitro and Palkovits, 1981;
handle' cell group with the substantia innominata proper (Figs
used for completely different structures. Authors often find it Olszewski and Baxter, 1954; Palkovits and Jacobowitz, 1974;
21-26, 80-84, 91-96) by Zaborszky. We had the choice of
necessary to include a table of equivalence of the different terms Pellegrino et al., 1979; Sidman et al., 1971; Wunscher et al.,
retaining the name substantia innominata for the rostral sliver
used for a given structure and such tables are becoming lengthier 1965). Our greatest debt is to the rat brain atlas of Konig and
only and calling the posterior part the sublenticular substantia
as new synonyms are added while the old ones refuse to die. To Klippel (1963).
innominata or calling all of this region substantia innominata;
quote Ann Graybiel, 'neuroscientists would rather swap We have not undertaken a comprehensive review of early
we have opted for using the traditional name for all this region.
toothbrushes than nomenclature.' Differences in nomenclature studies on the naming and delineation of structures; rather we
present a formidable communication barrier for neuroscientists. have taken the expedient course of detailed analysis of the Ventral pallidum (VP) and the medial forebrain bundle (mfb):
We believe that this atlas can contribute to the standardization current literature supported by extensive informal peer review of Nieuwenhuys et al. (1982) identified an 'a' and a 'b' component
of nomenclature; we therefore took the opportunity afforded by our proposed designations. We have relied heavily on the in the medial forebrain bundle. Rostrally, the 'a' component
the construction of the second edition to consult over 40 published historical surveys and on the encyclopedic knowledge occupies the dorsal part of the ventral pallidum (Figs 11-16).
neuroscientists on the appropriateness of our terminology. In and fine critical sense of our colleague. Bill Mehler. We have The 'b' component is found within the nucleus of the horizontal
accepting a particular term over other synonyms, we were included some notes on the basis of delineations of structures limb of the diagonal band (Figs 15-21, 79-81).
guided by the suggestions of our advisers and by the following (see below) in which we refer extensively to The Rat Nervous
Islands of Calleja (ICj): The Islands of Calleja were outlined
principles: (a) terms can be ratified by usage; (b) a viable System (Paxinos, 1985b) and in each section we cite relevant
according to Fallon et al. (1978).
'systematic' term is preferable to a slightly more frequently used review articles which support our choice of nomenclature and
but misleading synonym; (c) eponymous terms are to be delineations. Newly identified areas or designations that need Magnocellular preoptic nucleus (MCPO): In accordance with
dispensed with when viable alternatives exist. further elaboration than is provided in The Rat Nervous System other authors (e.g. Saper, 1984) we recognize a magnocellular
are dealt with individually in a paragraph or two. The preoptic nucleus lateral to the nucleus of the horizontal limb of
Abbreviations arrangement of headings corresponds to Paxinos (1985b). the diagonal band (Figs 18-23, 82, 83, 90, 91). These two nuclei
are poorly distinguished from each other (Zaborszky et al.,
The abbreviations used to label the drawings were constructed so Olfactory System 1986) and in the first edition of this atlas they were both
as to give a strong clue to the identity of the structures they considered to belong to the nucleus of the horizontal limb of the
represent. After considering the main features of the Refer to Switzer et al. (1985) for a general description of the diagonal band.
abbreviations used by the St Louis group (e.g. Saper and Loewy, olfactory system and de Olmos et al. (1985) for the olfactory
Basal nucleus (B): The cholinergic cells of the basal nucleus of
1980), Oswaldo-Cruz and Rocha-Miranda (1968) and other amygdala and the nucleus of the lateral olfactory tract. The
Meynert straddle the border between the globus pallidus and the
authors, we adopted the following guidelines for the authors review their own work in this field as well as the work of
internal capsule, invading both of these structures (Figs 21-30,
construction of our abbreviations. others (e.g. de Olmos et al., 1978; Haberly and Price, 1978).
82-86, 94-101). For a description of the distribution of
The abbreviations represent the order of words as spoken in We have reverted to the term 'piriform cortex' because the
cholinergic neurons in the basal forebrain see Sofroniew et al.
English (e.g. DLG = dorsal lateral geniculate nucleus). Capital functional term 'primary olfactory cortex' should logically
(1985) and Paxinos and Butcher (1985).
letters represent nuclei, and lower-case letters represent fiber include in it other olfactory recipient areas such as the olfactory
tracts. Thus the letter 'N' has not been used to denote nuclei, tubercle.
Septum, Hypothalamus, and the Neurosecretory
and the letter H' has not been used to denote fiber tracts. The
general principles used in the abbreviations of the names of Basal Ganglia and Basal Forebrain Nuclei
elements in the periodic table were followed as closely as
possible—the capital letter representing the first letter of a word Refer to Heimer et al. (1985) for a general description of the Refer to Bleier and Byne (1985) for a general description of the
in a nucleus is followed by the lower case letter most basal ganglia, and to Cuello et al. (1985) for the chemically septum and hypothalamus. In the first edition, the subdivisions
characteristic of that word (not necessarily the second letter: e.g. specified connections of the basal ganglia. Heimer et al. (1985) of the septum were based on the work of Swanson and Cowan
Mg = magnesium; Rt = reticular thalamic nucleus). review work in their laboratory that led to the identification of (1979), and the subdivisions of the hypothalamus were based on
XI