brain collections at the national museum of health and medicine

Ann. N.Y. Acad. Sci. ISSN 0077-8923

ANNALS OF THE NEW YORK ACADEMY OF SCIENCESIssue: Resources and Technological Advances for Studies of Neurobehavioral Evolution

Brain collections at the National Museum of Healthand Medicine

Archibald J. Fobbs, Jr.1 and John I. Johnson2

1National Museum of Health and Medicine, Armed Forces Institute of Pathology, Washington, DC. 2Radiology Department,Division of Anatomy, and Neuroscience Program, Michigan State University, East Lansing, Michigan

Address for correspondence: John I. Johnson, Radiology Department, Division of Anatomy, and Neuroscience Program,Michigan State University, A519 East Fee Hall, East Lansing, MI 48824-1316. [email protected]

Owing in large part to the foresight and efforts of Wally Welker, the National Museum of Health and Medicine hasbecome a major repository for collections of brain specimens vital to the study of neurobehavioral evolution. Fromits origins in the Armed Forces Institute of Pathology, with the collection of largely pathological specimens assembledby Paul Yakovlev, the museum has added to its resources four additional extensive collections, largely consisting ofspecimens acquired specifically for comparative and evolutionary studies: Welker’s collection from the University ofWisconsin-Madison, John I. Johnson’s collection from Michigan State University, the Adolf Meyer Collection fromthe Johns Hopkins University, and the Elizabeth Crosby collections from the University of Michigan. We describehere the history and contents of each of these five collections, to inform the scientific field of the extent and detailsof these remarkable resources.

Keywords: neuroanatomy; evolution of brains; comparative neuroscience; neurobiodiversity; brain databases

Introduction

Wally Welker, as he was nearing retirement, searchedfor a home for the extensive collection of brainsin the Physiology Department at the University ofWisconsin, including his own acquisitions, thoseassembled by Clinton Woolsey, and some speci-mens brought by Jerzy Rose from the Meyer PhippsCollection (see below) at Johns Hopkins Univer-sity, where this valuable material would be pro-tected from destruction consequent upon admin-istrative changes in missions and goals in universitystructure.

His quest found a like-minded response fromAdrianne Noe, in charge of collections at the Na-tional Museum of Health and Medicine (NMHM)

The opinions or assertions presented hereafter are theprivate views of the authors and should not be construedas official or reflecting the views of the Department ofDefense, the National Museum of Health and Medicine,the Armed Forces Institute of Pathology, or the U.S. ArmyMedical Readiness and Materiel Command.

at the Armed Forces Institute of Pathology (AFIP),located in the Walter Reed Army Medical Center inWashington, DC. The NMHM was the major repos-itory of collections of embryos from major sourcesaround the country and also possessed the greatYakovlev-Haleem brain collection (see below). Noeand Welker saw the addition of the Wisconsin col-lection to the Yakovlev-Haleem as a first step informing a similar central national home for braincollections.

Welker secured significant assistance in identify-ing sources of support, from Christopher Platt, thenat NSF, who also saw the great advantage of bring-ing collections together at a single location. Thisconsolidation greatly facilitates broad studies by re-searchers, and aids teachers seeking instructionalmaterials, in a sanctuary protected from the vagariesof scientific ebbs and flows in current topical mate-rials. Scattered university facilities are necessarily atrisk, and many specimens and collections have beenlost to these winds of change. This joint collectionprinciple attracted more additions to the criticalmass: brains from Michigan State University (see

doi: 10.1111/j.1749-6632.2011.06036.xE20 Ann. N.Y. Acad. Sci. 1225 S1 (2011) E20–E29 c© 2011 New York Academy of Sciences.

Fobbs & Johnson Brain collections, National Museum Health Med.

below)—an adjunct of the Wisconsin collection—the great and historic Crosby Collections (see below)at the University of Michigan (not to be confusedwith Michigan State University), from the MeyerPhipps Collection—the ancestor of UW and MSUcollections—and other smaller groupings, includ-ing the Pubols Collection from Oregon Health Sci-ences University, the Lockard Collection from theMedical University of South Carolina, and the StarrCollection from Columbia University.

Financial support for the transfer of the Wis-consin, Michigan State, and Michigan collectionswas provided by the National Science Foundation,along with other sources that assisted in the gen-eral process of collection amalgamation. The valueof the brain and embryo collections was recog-nized in the federal law governing the closure ofthe Walter Reed Campus in assuring the protec-tion and continuation of the NMHM as a nationalresource.

In the tables of contents of each of the collectionspresented below, we have preserved, as far as possi-ble, the format and nomenclature used by the orig-inal curators of the collections. To alter and makeassumptions about what is meant by entries in thecatalogs risks distortions and errors; users of thecollections will be better able to judge what con-temporary nomenclatures and classifications will beappropriate for their purposes.

The Yakovlev-Haleem Collection

History of the collectionPaul Yakovlev was born in 1894 and received hisdegree in medicine from the Military MedicalAcademy in Leningrad in 1919. He left Russia laterthat year, walking across the frozen Gulf of Finlandwith only a satchel and his medical diploma.1

Eventually, he migrated to Paris where he stud-ied under the famous neurologists Pierre Marie andF. J. Babinski. He received an M.D. degree fromthe University of Paris in 1925. In the same year,he emigrated to the United States and practicedmedicine in Providence, Rhode Island. He met Stan-ley Cobb and William G. Lennox of Harvard Uni-versity and worked under them in the neurologyclinic of Boston City Hospital. He became the liai-son between Harvard and the Monson State Hospi-tal at Palmer, Massachusetts. According to Yakovlev,“that’s where I began my career as a scholar.”1

Lennox had wanted him “to make some sense” out

of the epileptic patients at the hospital and their dis-order. In contrast to his vocation of neurologist tothe patients, “by avocation” as Yakovlev put it, hebegan to study by autopsy some of the brains of hisformer patients.

Lacking proper facilities and techniques forproper preparation of autopsy material, Yakovlevsecured a fellowship to study in Switzerland at Con-stantin von Monakow’s Hirnanatomisches Institutin Zurich, where he studied with M. A. Minkowski.At the institute he vividly recalled long lines of cab-inets containing serial sections with “some millionsof slides.” Upon return to Monson he looked into theconstruction of a proper microtome to adequatelyprepare sectioned brains for study, as well as stain-ing and mounting techniques, and thus was bornthe eventual great Yakovlev Collection of normaland pathological human brains. He continued thisenterprise in his later assignments.

In 1936, Yakovlev became director of laborato-ries at the Metropolitan State Hospital in Waltham,Massachusetts. In 1938, he became clinical direc-tor of research laboratories at the Walter E. FernaldSchool in Waltham, Massachusetts, where he pre-pared many of the specimens in this famous col-lection of serial histological sections of over 1,000brains. All the slides (over 250,000) in this collectionwere carefully prepared, sectioned and stained by thesame exacting technique. The collection moved tothe Warren Museum of Harvard University whereYakovlev became associate clinical professor of neu-ropathology and curator of the Warren AnatomicalMuseum.

After his retirement, the collection moved backto the Fernald School. In 1974, by a joint agreementwith Yakovlev, Harvard University, the National In-stitutes of Health and the AFIP, this outstanding col-lection was transferred to the AFIP and its NMHMin Washington, DC, where it is now available as a na-tional resource for the research studies by qualifiedscientists.

In addition to the specimens acquired person-ally by Yakovlev, many specimens were donatedto the collection from a great variety of sources.For comparative and evolutionary research, notablecontributions include monkey brains from Mor-timer Mishkin, of the National Institute of Men-tal Health, and from Samuel Gelfan and others atthe Yale Aeromedical Research Unit at Yale Uni-versity; as well as dolphin brains donated by Peter

Ann. N.Y. Acad. Sci. 1225 S1 (2011) E20–E29 c© 2011 New York Academy of Sciences. E21

Brain collections, National Museum Health Med. Fobbs & Johnson

Figure 1. Illustrations of stained sections in the Yakovlev-Haleem Collection. Images A and B show sections in the horizontalplane, C and D in the parasagittal plane. Stains used with these sections are cresyl violet, Nissl method in A and C, and ironhematoxylin, Loyez version of the Weigert method in B and D. Fully labeled copies of these images can be seen in online atlases athttp://www.brains.rad.msu.edu, or https://www.msu.edu/∼brains//brains/index.html.

Morgane from the Communications Research Insti-tute in Miami, Florida.

Mohammed Haleem was associated withYakovlev’s brain collection for more than thirtyyears. He was born in Palestine, and came tothe United States in 1960, settling in Boston,Massachusetts where he first began working withYakovlev. Haleem came to the AFIP in 1974 whenYakovlev relocated the collection to AFIP from Har-vard Medical School. In 1983, Haleem became cu-rator of the Yakovlev Collection, which, in honorof his dedication and contributions, was renamedthe Yakovlev-Haleem Collection in February 1994.Haleem is remembered today for his unmatchedskills in preparing brain slices for study.

Contents of the collectionMost of the brains in the collection were embed-ded in celloidin, and were sectioned at intervals of

35 �m. The stained sections are mounted in pairs ofstained sections at intervals of from 5 to 25 sections.One member of each pair is stained with cresyl vio-let by the Nissl method to show cell bodies; the otheris stained with iron hematoxylin by the Loyez mod-ification of the Weigert method to show myelinatedfibers (Fig. 1). Many of the intervening, unstained,sections are preserved in individual containers forpossible future use.

The collection is catalogued here in two parts.The Human Collection (Table S1A) includes 433human brains specimens designated as “normativecontrols” in what constitutes a developmental series,ranging in age from embryos to the 10th decade.The Comparative Collection (Table S1B) consistsof brains of three cats, 3 embryonic fur seals, 20embryonic and adult whales and dolphins, 1 horse,38 chimpanzees, 1 gorilla, and 97 fetal and adultrhesus monkeys.

E22 Ann. N.Y. Acad. Sci. 1225 S1 (2011) E20–E29 c© 2011 New York Academy of Sciences.


In addition to the specimens listed here, theYakovlev-Haleem assembly also includes another576 specimens of a great variety of pathologi-cal conditions, including 129 malformations, 149instances of miscellaneous neuropathology, 49cases of cerebrovascular strokes, and 92 resultsof neurosurgery, as classified by Kretschmannet al.2

The Welker Wisconsin Collection

History of the collectionThis history is based on that compiled by WallyWelker and published on the website http://brainmuseum.org.

The Welker Collection was started in the 1950s byClinton Woolsey. Woolsey came from Johns Hop-kins to start brain research at the University ofWisconsin–Madison, soon after the Second WorldWar. He was the founder of the laboratory, and laterthe Department of Neurophysiology at Wisconsin.He brought with him not only a strong and extensiveinterest in the comparative anatomy of mammalianbrains, but also the beginnings of what has becomeone of the premier collections of its kind anywherein the world.

Konrad Akert played a role in standardizing tech-niques of perfusion and histological processing andprotocol keeping in the early years. The brain collec-tion was increased and diversified by Wally Welkerand colleagues over the next four decades. Dur-ing this period, a large number of faculty andstudents carried out comparative neurophysiolog-ical and neuroanatomical analyses in a variety ofmammals.

Many of these studies were initiated because ofthe special behavioral repertory of the different an-imals studied. John I. Johnson joined these investi-gators for postdoctoral training in 1960. By the timeJohnson arrived, Welker, a member of the Wiscon-sin faculty, had begun the formal collection of brainspecimens with five-year support from the NationalInstitutes of Health for “comparative study of brainmorphology in mammals.”

With the advice of Konrad Akert and Jerzy Roseat Wisconsin, together with the collaboration of nu-merous students and colleagues, Welker assembledbrains from a wide variety of mammals from a repre-sentative variety of mammalian orders—specimensthat were perfused and prepared by standardizedhistological procedures in ways that would provide

neuroanatomical study materials for many genera-tions of comparative neuroanatomists.

Welker worked together on these comparativeprojects with Johnson over the last several decadesand both worked together with Jon Kaas, John All-man, Michael Merzenich, and many others over thistime span. The neuroanatomical material that wasassembled was transported and is now curated andmaintained at the NMHM in Washington, DC, un-der the supervision of the museum’s director, Adri-anne Noe, and curator, Archibald Fobbs, and ismade widely available for study by students, edu-cators, and researchers worldwide.

Contents of the collectionThe Welker Wisconsin Collection consists in a to-tal of 409 specimens with representatives of 122species. Most of these are mammalian brains. A fewspecimens of brains from other vertebrate cladesare included: amphibian salamander and bullfrogs;reptilian alligators, Tegu lizards, iguana, Westernrattlesnake and turtles; and avian ostrich, owls, gulls,and pigeons (Table S2). Some special features are adevelopmental series of brains of rats and cats and anextensive experimental series of brains of echidnas,galagoes, squirrel monkeys, chimpanzees, beavers,mice, rats, rabbits, cats, raccoons, and manatees.

As collecting proceeded through the years, a stan-dard protocol was gradually established, includingmetric photography of the living animal as well as offixed specimens of bodies and brains, and the man-ufacture of acrylic casts of the brain of one specimenfrom each species.

The brains were then embedded in celloidin, or insome special cases in paraffin, and were sectionedat intervals of 15–60 �m, depending on the sizeof the brain. The stained sections are mounted inpairs of stained sections at intervals of from 1 to10 or more sections, again depending on brain size.One member of each pair is stained with thioninby the Nissl method to show cell bodies; the otheris stained with iron hematoxylin by the Weil, Ma-hon, or Sanides-Heidenhain-Woelcke3 modifica-tions of the Weigert method to show myelinatedfibers (Fig. 2). The sectioning and staining, as seenin this figure, was done in accordance with veryexacting standards by a team of long-working his-tological technicians, supervised for many decadesby Helen Brandemuehl, who had been person-ally trained by Cecilia Bisson, Adolf Meyer’s chief



Figure 2. Illustrations of stained sections in the Welker Wisconsin Collection. Sections are in the coronal plane. Images A and Care stained with thionin, Nissl method; B and D are stained with iron hematoxylin, the Sanides-Heidenhain-Woelcke modificationof the Weigert method. Additional illustrations of specimens in this collection can be viewed at http://brainmuseum.org.

histologist at the Hopkins Phipps Clinic as arrangedby Clinton Woolsey when he founded the labora-tory; in later years the Bisson-Brandemuehl tradi-tion was continued by Inge Siggelkow.

Johnson Michigan State Collection

History and contents of the collectionThe arrival in the Wisconsin laboratory of a zoospecimen of an Australian Western Gray Kanga-roo, in 1962, highlighted a notable gap in the Wis-consin collection. There was a dearth of specimensfrom the great marsupial branch of the mammalianradiation. John I. Johnson was stimulated by thisdiscovery to plan an expedition to Australia to fill

this gap, by means of a Fulbright fellowship andgrant support from the National Institute of MentalHealth. With much assistance from Peter Bishop, hisAustralian host, at that time professor and chair ofphysiology at the University of Sydney, and MichaelMarsh in the Sydney Zoology Department, he wasable to obtain specimens of many local marsu-pial species. In addition brains were obtained fromspecies of Catshark and avian Emu from experi-ments in cardiovascular physiology in the Sydneydepartment.

Marsh introduced Johnson to John A. W. Kirsch, agraduate student at the University of Western Aus-tralia who was in Sydney to obtain specimens of



various tissues from eastern Australian species forhis dissertation research, with W. D. L. Ride at theUniversity of Western Australia, on marsupial phy-logeny as revealed in comparative serology. Throughthis connection, and the assistance of Peter Bishop,he was able to travel to Perth, where with Kirsch’s as-sistance specimens were obtained from the coloniesmaintained in the UWA Zoology department. LaterKirsch and Johnson jointly toured Tasmania, withexcellent assistance from the Animals and Birds pro-tection board of Tasmania, and were able to obtainfrom their native habitats specimens of TasmanianDevil (now in danger of imminent extinction), Na-tive Cat, Tasmanian Pademelon, Rufous Wallaby,Potoroo, and a monotreme Echidna.

The Johnson and Kirsch collaboration contin-ued during Kirsch’s postdoctoral fellowship at theUniversity of Kansas, which included an expedi-tion through South America gathering specimensof the South American marsupial radiation. Whileobtaining his various tissue and serological spec-imens for further work in marsupial phylogeny,Kirsch, after his experience in Australia, was ableto also perfuse brains for addition to the brain col-lections, including such phylogenetically significantand rare specimens as Caenolestes obscurus, Lestorosinca, and Dromiciops australis. Additional SouthAmerican species, including marsupials Didelphisalbiventris, Marmosa murina and Thylamys veluti-nus, were kindly donated by Eduardo Oswaldo-Cruzand Carlos Rocha-Miranda at the Federal Univer-sity of Rio de Janeiro, as were specimens of placentalagoutis by Drs. Angelo Machado and Gilberto Cam-pos at The Federal University of Minas Gerais in BeloHorizonte.

Another significant gift to the collections was acomplete set of “extra” unstained sections of brainsof several species of bats, mostly Caribbean, by Dr.Paul Pirlot of the Universite de Montreal. Thesesections, embedded in paraffin, were subsequentlymounted and stained at Michigan State. They arelikely to be the only survivors of the large Pirlotcollection; the rest of the collection was destroyedin a disastrous fire in Quebec.

The Michigan State collection also includes nor-mal specimens of several species maintained atthe university for experimental research, includingAnolis lizards, bobwhite quail, sheep from the en-docrine research unit in physiology, kangaroo rats,various species of Peromyscus deer mice from var-

ious habitats from the colony of John A. King inzoology, as well as local opossums, fox squirrels,and raccoons from experiments in the Johnsonlaboratory.

A complete listing of specimens in the collectionincludes 81 species, represented by 177 specimens,83 placentals, 13 bats, 69 marsupials, 5 monotremes,and 19 non-mammals (Table S3).

Processing of all these specimens followed, as faras was possible, the protocols and procedures usedat Wisconsin by Wally Welker, including photog-raphy of the living animal as well as metric pho-tographs of fixed specimens of bodies and brains;manufacture of acrylic casts of the brain of one spec-imen from each species, embedding in celloidin andstaining with alternate sections series stained withthionin by the Nissl method, and with hematoxylinto show myelinated fibers, using either the Weil orthe Sanides-Heidenhain-Woelcke3 variants of theWeigert technique (Fig. 3). Sectioning, staining, andmounting of the stained sections on glass slideswas the work of many skilled technicians and stu-dents over the decades, with notable contributionsby Grace Heung-Sook Chung, Tony Duke, San-ford Feldman, Emmeline Gatewood Haight, JohnR. Haight, Paul Herron, Joanna M. Hill, MichaelOstapoff, Michael Peterson, Edwin Rubel, SharleenSakai, Steven Warach. Michael Widener, Robert C.Switzer III, Philip Ulinski, W. Lee Weller, and SidneyWiener.

Meyer Phipps Hopkins Collection

History of the collectionIn 1908 the Philadelphia steel magnate HenryPhipps visited the tuberculosis clinic he had fundedat the Johns Hopkins University, and asked if therewere another opportunity for his philanthropy.Dean William Henry Welch of the Hopkins Med-ical Faculty took the opportunity to found a de-partment of psychiatry, in a state-of-the-art clinicfunded by Phipps, and hired Adolf Meyer to over-see the foundation of the clinic, department, andcurriculum. Meyer was Swiss, and professor of psy-chiatry at Cornell University at that time; he believedin a scientific basis for Psychiatry, emphasizing bi-ological factors and life experiences as explanatory,preferring data-based to theoretical formulations.He originated the terms and the fields of biopsychol-ogy, and of mental hygiene. This thinking impressedDean Welch, and this was his basis for recruiting



Figure 3. Illustrations of stained sections in the Johnson Michigan State Collection. Images A and C are stained withthionin, Nissl method; B and D are stained with iron hematoxylin, B using the Weil modification of the Weigert method,and D the Sanides-Heidenhain-Woelcke modification. Additional illustrations of specimens in this collection can be viewed athttp://www.brains.rad.msu.edu or https://www.msu.edu/∼brains//brains/index.html. Under Title 17 of the U. S. Code, Section 105,copyright protection is not available for any work of the United States Government. The opinions or the assertions herein are theprivate views of the authors and are not to be construed as official or as reflecting the views of the National Museum of Health andMedicine, the Department of the Army, or the Department of Defense.

Meyer to oversee the development of psychiatry atHopkins.4

Accordingly the clinic at Hopkins would havepatients, laboratories, and experimental and ob-servational research, including extensive case histo-ries and anatomical and physiological laboratories.Meyer had, in his own training in Switzerland, stud-ied with the renowned neuroanatomists AugusteForel and Constantin von Monakow (at the sameinstitute where Paul Yakovlev learned about braincollecting in subsequent years). As a result, Meyer,like Yakovlev after him, believed in having adequateanatomical and physiological tools for psychiatricresearch. This included an ever-growing collectionof neuroanatomical material, including brains ofanimals used in experimental research.

The neuroanatomical material was prepared us-ing the best technology and standards, and inMeyer’s long tenure this resulted in the exten-sive brain collection described here. Many classical

anatomical studies resulted from use of these mate-rials beyond the reasons for their initial collection.For example, the sheep brains that were collectedand processed by Charles Bagley, in the course ofexperimental physiological studies,6 were later usedby Jerzy Rose for his classical descriptions of thala-mic and cerebral cortical architecture.7,8 When JohnI. Johnson began neuroanatomical study, his first in-struction from Wally Welker was to learn the organi-zation of the sheep thalamus as described by Rose,8

and this particular exercise is still a most instructiveand thorough introduction to the architecture of themammalian thalamus.

In several reorganizations of neuroscience atHopkins in the years after the Meyer era, VernonMountcastle, who studied with Meyer as a medicalstudent, stayed at Hopkins his entire professionallife, became chairman of the Hopkins physiologydepartment, founder of the Hopkins neuroscienceprogram, and was the first president of the



Society for Neuroscience. Seeing the value and theneed for preservation of the priceless Meyer Col-lection, Mountcastle was instrumental in arrangingits transfer to NMHM to join the Wisconsin andMichigan State Collections; themselves direct de-scendants, in outlook, methodology and organiza-tion, of the Meyer collection.

Unlike these descendant collections, the MeyerPhipps Collection was not designed to be a phylo-genetic or zoological collection. This is reflected inthe highly general naming of species and clades ofthe animals whose brains were collected originallyfor various experimental purposes. However, thisdefect as a systematic biological collection is offsetto a degree by the fact that there exists a wealth ofexperimental articles containing physiological andbehavioral data directly related to the neuroanatomyin the stained sections.

Contents of the collectionThe largest component of the 267 specimens in thecollection is a group of 197 mammal brains belong-ing to 56 species. Other holdings are 22 bird brainsfrom 13 species, 34 brains of 11 species of reptiles,6 brains of one amphibian species, 17 brains of 5species of bony fish, and one shark brain (Table S4).The generally well-stained sections are embeddedin celloidin: 169 specimens are stained for cells, 75other specimens show myelinated fibers, and 15 havealternating fiber and cell-stained sections.

The Huber-Crosby and Crosby-LauerCollections

History of the collectionsThe assembly of the Yakovlev, Wisconsin, andMichigan State collections at the NMHM attractedthe attention of Sarah Winans Newman, chairper-son of the Department of Anatomy and Cell Biologyat the University of Michigan at Ann Arbor. The de-partment had custody of the extensive assemblies ofover 1,000 brain specimens collected by ElizabethC. Crosby, working with G. Carl Huber until hispassing, and with her profusion of students duringher six decades of teaching and researching in hu-man and comparative vertebrate neuroanatomy inAnn Arbor. Newman had concerns about the even-tual fate of these extraordinary resources. With hersupport and the copious assistance of the curator ofthese materials, Beverly Shepard, this accumulation

of specimens of vertebrate brains was cataloguedand transferred to the NMHM.

The collection is divided into two parts, an earlierHuber-Crosby and a later Crosby-Lauer Collection.Huber was the anatomical laboratory director whohired, collaborated with, and supported Crosby formore than two decades. Edward W. Lauer was aCrosby student who served many years as a facultymember in the department.

The Huber-Crosby Collection, along with that ofC. U. Ariens Kappers in Amsterdam (see the articleby Hofman in this volume), formed the foundationof what became “The Comparative Anatomy of theNervous System of Vertebrates, Including Man,”9

the three volume standard and encyclopedic refer-ence for vertebrate neuroanatomy for over half acentury, from the 1930s to the 1990s. The authorsare listed as Ariens Kappers, Huber and Crosby, butit is widely recognized that this listing should bein the reverse order, as is the case with the Huber-Crosby Collection. Crosby was deferential to Huberin memorial respect for his support and collabora-tion over the years, and listed his name before hersin both the book and in the naming of the collec-tion. Crosby spent some time in Amsterdam withC. U. Ariens Kappers, and was invited by him toundertake the translation,10 which soon became therevision and updating, of his earlier massive work,Die vergleichende Anatomie des Nervensystems derWirbeltiere und des Menschen.11,12

The later Crosby-Lauer Collection was largely as-sembled by Lauer, whom Crosby had encouragedto come to graduate school; he eventually becameher fellow faculty member and continuing collab-orator. This collection contains representatives ofspecies not included in the Huber-Crosby Collec-tion. A beautifully stained specimen of the brain ofa greater panda, Ailuropoda melanoleuca. is listed inboth collections; it was a resource used by Lauer inhis study of the olfactory system in this species.13

Contents of the collectionsThe Huber-Crosby Collection, (Table S5A) alongwith that of C. U. Ariens Kappers in Amsterdam,contains specimens of 15 cyclostomes, 5 sharks, 17bony fish, 47 amphibians, 122 reptiles, 131 birds and393 mammals, making a total of 730.

The Crosby-Lauer Collection (Table S5B) in-cludes specimens of 2 cyclostomes, 5 bony fish, 5amphibians, 7 reptiles, 14 birds, and 58 mammals.



Discussion

Problematic species identificationsFor the listings of contents of all of the collections, wepresent the names as chosen, listed, and organizedby the donors or curators of the collections at thetime of accession. Many of the names are currentlyconsidered obsolete; however, obsolete names areoften resurrected during long useful lives that canextend over decades and into centuries. More prob-lematic are vague designations, such as “snake,” butit is dangerous to guess at the true identity of a spec-imen. It is conceivable that in the not too distantfuture, it will be possible to more securely identifyspecimens through DNA analysis of a bit of tissue.This is one importance of preserving actual tissuesrather than only digital representations.

The importance of digitized renditionsof collected specimensHowever, there is no reason to demean the tremen-dous value of digital representations of collectionspecimens. The article by Jones, Stone, and Karten,in this volume, makes an excellent case for the valueof modern imaging methods in making availableinformation, resident in collection specimens, towidespread and varied audiences.

Wally Welker was a pioneer in using digitalphotography and the Internet in making availableknowledge about the specimens he assembled, onthe website http://brainmuseum.org. The brain mu-seum images have become a standard source for ed-ucational and general publication purposes, whenpictures of whole brains or sample sections areneeded. We are making use of several of them inthis paper to show some of the value of this material(Figs. 1–3). The article by Buchanan and Johnsonin this volume illustrates how they can be used toadvance comparative neuroscience.

According to Google statistics, The Human BrainAtlas at https://www.msu.edu/user/brains// is themost widely used online human brain atlas, anddata retrieved from permission requests show thatit is used at all educational levels on all continents,as well as in textbooks, research publications andthe popular press. The atlas images are from theYakovlev-Haleem Collection, an interesting use, forgeneral information, of material originally assem-bled for pathological diagnoses and histories. Thecompanion Sheep Brain Atlas at this site is in similarwidespread use in veterinary education and for gen-

eral neuroscience laboratories where cheap sheepbrains are the demonstration materials in commonuse. Images in the Sheep Brain Atlas are from theWelker Wisconsin and Johnson Michigan State col-lections. A unique Dolphin Brain Atlas at the samelocation has images of sagittal and horizontal sec-tions from the Yakovlev-Haleem Collection; a seriesof images of coronal sections from the Welker Wis-consin Collection is in preparation for addition tothe atlas.

These are but initial examples of the momentousimpact brain collection material, in digital tran-scription, can have on all branches of human in-formational activities worldwide. Digital renditionof collection resources is a vital activity, and canbe pursued as needed for specific purposes. Adop-tion of standard digitizing formats and methodscan result, over long periods of time, in formidabledatabases assembled from individual digitizingprojects.

Conflicts of interest

The authors declare no conflicts of interest.

Supporting Information

Additional supporting information may be foundin the online version of this article:

Table S1: Detailed lists of specimens in theYakovlev-Haleem Collection, useful for studies ofvertebrate brain evolution. (A) List of normativecontrol human specimens in sequence by age. (B)List of nonhuman vertebrate specimens.

Table S2: List of specimens in the Welker Wiscon-sin Collection, arranged alphabetically by commonnames of species. The specimen numbers designatethe year of acquisition by the first two digits, fol-lowed by a hyphen and 1 to 3 digits denoting theorder in which the specimens were acquired. Thus63–361 designates the 361st specimen acquiredin the Wisconsin laboratories in the year 1963.The stain listed as Heidenhain is the Sanides-Heidenhain-Woelcke modification of the Weigertmethod, introduced to the Wisconsin laboratoriesby Friedrich Sanides in the mid-1960s.

Table S3: List of specimens in the Johnson Michi-gan State Collection, arranged in approximate phy-logenetic order, according to the Universal SpeciesIdentification System.5 The specimen numbers for



the Michigan State Collection carry the followinginformation. The first two digits represent the yearof acquisition. The third digit gives some informa-tion about the purpose of acquring the specimen –0 is a normal marsupial or monotreme specimen,1 is an experimental specimen, 2 is a normal spec-imen, 3 is a cat, 4 is an opossum, and 5 is a rac-coon. The last two digits represent the sequence dur-ing that year in which the specimen was acquired.Thus 65063 indicates a normal monotreme or mar-supial specimen, the 63rd specimen acquired in1965.

Table S4: List of specimens in the Meyer PhippsJohns Hopkins Collection.

Table S5: Lists of specimens in the Huber-Crosbyand Crosby-Lauer collections.

Please note: Wiley-Blackwell is not responsiblefor the content or functionality of any sup-porting materials supplied by the authors. Anyqueries (other than missing material) shouldbe directed to the corresponding author of thearticle.

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brain collections at the national museum of health and medicine

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