5
A Tribute to Berta Scharrer STEVEN L. WISSIG* Department of Anatomy, University of California, San Francisco, California During my graduate studies at Yale University al- most 50 years ago, Sanford Palay, a newly appointed instructor in the Department of Anatomy, was my thesis adviser. As you will read below, earlier during his medical training he had worked in the research labora- tory of Ernst and Berta Scharrer at Western Reserve University. Thus he had become familiar with the studies they had carried out on neurosecretion prior to 1950, and he was beginning a research project in the same area at Yale. Although my own thesis project was in a different area, we had many discussions about the possible physiologic significance of the neurosecretory system, which presaged the explosion of information about the link between the nervous and endocrine sys- tems that would emerge during the next two decades. I met Berta Scharrer for the first time in the Depart- ment of Anatomy at the Albert Einstein College of Medicine in the late seventies when I spent part of a sabbatical leave working there with Birgit Satir. Berta and I became close friends and had many conversations then and during my subsequent visits to Einstein. Her sharp mind, devotion to science, and warm personality made a strong impression on me. When she died and I was asked to write her biography for the Anatomists, I willingly accepted the invitation in order to provide members of the Association, who had not had personal contact with her, a fuller insight into her life and accomplishments. Berta Vogel Scharrer was born December 8, 1906 in Munich, Germany, into a prosperous, well-educated family. Her father was a judge serving as vice president of the Federal Court of Bavaria. As a young student she became interested in biology, and she had already made an important decision concerning her future career even before she entered the University of Munich. She would study to become a research scientist in biology, although she knew that the chances of a woman achieving this goal in Germany were virtually nil. She had the good fortune to have, as her thesis adviser at the university, Karl von Frisch who later earned a Nobel Prize for his behavioral studies of the honeybee. For her own research project she compared the taste and nutritive value of various sugars for this insect. At the university she met a fellow student, Ernst Scharrer, whom she would wed in 1934, four years after receiving her doctorate. After her graduation she and Ernst worked at the Research Institute of Psychiatry in Munich where she studied spirochaete infections of the brain in birds and amphibians. When Ernst received an appointment at the Edinger Institute of Neurology in Frankfurt-am- Main in 1934, Berta accompanied him, working as an unpaid research associate. At this stage of their careers, the Scharrers spent summers at the Zoological Station in the Bay of Naples. Ernst had earlier, in 1928, published a paper describing the presence of secretory droplets in certain hypotha- lamic neurons of a marine teleost. His intriguing pro- posal that nerve cells were capable of secreting mate- rial as well as conducting impulses was greeted with scepticism at best. Meanwhile, Berta turned her atten- tion to the nervous system of invertebrates and uncov- ered morphologic evidence of neurosecretory activity in an opisthobranch mollusc (1935) and a marine worm (1936). At this point the Scharrers had established the pattern that guided their scientific careers until the time of Ernst’s untimely death in 1965. Their research would center on an exploration of the significance of neurosecretion, Ernst examining the phenomenon in vertebrates, Berta in invertebrates. Where they worked would be determined by where Ernst accepted an academic position, and Berta would ‘‘tag along,’’ as it were, as an unpaid research associate. Only in this way did Ernst feel they could work in the same institution without being accused of nepotism. The Scharrers left Germany in 1937 as a matter of conscience; they were unable to countenance the poli- cies of the National Socialist state. To remain in Ger- many, Ernst, as a physician, would have had to accept a military appointment, and, as a scientist, he would have had to eschew and denounce his Jewish col- leagues. Worst still, he was repelled by the thought of being recruited for a position from which a Jewish colleague had been dismissed. They came to the United States where Ernst had obtained a one-year Rockefeller Fellowship in the Department of Anatomy at the Univer- sity of Chicago. Berta continued her work on inverte- brates initially with Drosophila, but subsequently and for the remainder of her research career with the cockroaches and their near cousins. In the United States, they made a series of moves dictated by the progress of Ernst’s academic career. Their year in Chicago was followed by two years in the laboratory of Herbert Gasser at the Rockefeller Insti- tute in New York City. Their work progressed so fruitfully that they could present a landmark paper on neurosecretion at the annual meeting of the Association for Research in Nervous and Mental Diseases in 1940, an event that introduced the neuroscience community to the concept of neurosecretion. That year the Schar- rers moved to the Department of Anatomy at Western Reserve University when Ernst was appointed assis- tant professor by the new chairman, Normand Hoerr, their close friend from the University of Chicago. The Scharrers remained at Western Reserve until 1946. *Correspondence to Steven L. Wissig, Dept. of Anatomy, University of California, San Francisco, CA 94143-0452. THE ANATOMICAL RECORD 249:1–5 (1997) r 1997 WILEY-LISS, INC.

A tribute to Berta Scharrer

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A Tribute to Berta Scharrer

STEVEN L. WISSIG*Department of Anatomy, University of California, San Francisco, California

During my graduate studies at Yale University al-most 50 years ago, Sanford Palay, a newly appointedinstructor in the Department of Anatomy, was mythesis adviser. As you will read below, earlier during hismedical training he had worked in the research labora-tory of Ernst and Berta Scharrer at Western ReserveUniversity. Thus he had become familiar with thestudies they had carried out on neurosecretion prior to1950, and he was beginning a research project in thesame area at Yale. Although my own thesis project wasin a different area, we had many discussions about thepossible physiologic significance of the neurosecretorysystem, which presaged the explosion of informationabout the link between the nervous and endocrine sys-tems that would emerge during the next two decades.

I met Berta Scharrer for the first time in the Depart-ment of Anatomy at the Albert Einstein College ofMedicine in the late seventies when I spent part of asabbatical leave working there with Birgit Satir. Bertaand I became close friends and had many conversationsthen and during my subsequent visits to Einstein. Hersharp mind, devotion to science, and warm personalitymade a strong impression on me. When she died and Iwas asked to write her biography for the Anatomists, Iwillingly accepted the invitation in order to providemembers of the Association, who had not had personalcontact with her, a fuller insight into her life andaccomplishments.

Berta Vogel Scharrer was born December 8, 1906 inMunich, Germany, into a prosperous, well-educatedfamily. Her father was a judge serving as vice presidentof the Federal Court of Bavaria. As a young student shebecame interested in biology, and she had already madean important decision concerning her future careereven before she entered the University of Munich. Shewould study to become a research scientist in biology,although she knew that the chances of a womanachieving this goal in Germany were virtually nil. Shehad the good fortune to have, as her thesis adviser atthe university, Karl von Frisch who later earned aNobel Prize for his behavioral studies of the honeybee.For her own research project she compared the tasteand nutritive value of various sugars for this insect. Atthe university she met a fellow student, Ernst Scharrer,whom she would wed in 1934, four years after receivingher doctorate.

After her graduation she and Ernst worked at theResearch Institute of Psychiatry in Munich where shestudied spirochaete infections of the brain in birds andamphibians. When Ernst received an appointment atthe Edinger Institute of Neurology in Frankfurt-am-Main in 1934, Berta accompanied him, working as anunpaid research associate.

At this stage of their careers, the Scharrers spentsummers at the Zoological Station in the Bay of Naples.

Ernst had earlier, in 1928, published a paper describingthe presence of secretory droplets in certain hypotha-lamic neurons of a marine teleost. His intriguing pro-posal that nerve cells were capable of secreting mate-rial as well as conducting impulses was greeted withscepticism at best. Meanwhile, Berta turned her atten-tion to the nervous system of invertebrates and uncov-ered morphologic evidence of neurosecretory activity inan opisthobranch mollusc (1935) and a marine worm(1936).

At this point the Scharrers had established thepattern that guided their scientific careers until thetime of Ernst’s untimely death in 1965. Their researchwould center on an exploration of the significance ofneurosecretion, Ernst examining the phenomenon invertebrates, Berta in invertebrates. Where they workedwould be determined by where Ernst accepted anacademic position, and Berta would ‘‘tag along,’’ as itwere, as an unpaid research associate. Only in this waydid Ernst feel they could work in the same institutionwithout being accused of nepotism.

The Scharrers left Germany in 1937 as a matter ofconscience; they were unable to countenance the poli-cies of the National Socialist state. To remain in Ger-many, Ernst, as a physician, would have had to accept amilitary appointment, and, as a scientist, he wouldhave had to eschew and denounce his Jewish col-leagues. Worst still, he was repelled by the thought ofbeing recruited for a position from which a Jewishcolleague had been dismissed. They came to the UnitedStates where Ernst had obtained a one-year RockefellerFellowship in the Department ofAnatomy at the Univer-sity of Chicago. Berta continued her work on inverte-brates initially with Drosophila, but subsequently andfor the remainder of her research career with thecockroaches and their near cousins.

In the United States, they made a series of movesdictated by the progress of Ernst’s academic career.Their year in Chicago was followed by two years in thelaboratory of Herbert Gasser at the Rockefeller Insti-tute in New York City. Their work progressed sofruitfully that they could present a landmark paper onneurosecretion at the annual meeting of the Associationfor Research in Nervous and Mental Diseases in 1940,an event that introduced the neuroscience communityto the concept of neurosecretion. That year the Schar-rers moved to the Department of Anatomy at WesternReserve University when Ernst was appointed assis-tant professor by the new chairman, Normand Hoerr,their close friend from the University of Chicago. TheScharrers remained at Western Reserve until 1946.

*Correspondence to Steven L. Wissig, Dept. of Anatomy, Universityof California, San Francisco, CA 94143-0452.

THE ANATOMICAL RECORD 249:1–5 (1997)

r 1997 WILEY-LISS, INC.

During this time Berta established a breeding colony ofthe South American woodroach, Leucophaea maderae(Orthoptera) and began a series of experiments inwhich she used surgical manipulations to define therole of the neurosecretory organs, i.e., the brain-corpuscardiacum-corpus allatum system, in the endocrinologyand development of insects. Another feature of theScharrers’ stay in Cleveland was their close associationwith Sanford Palay who began his training at theMedical School in 1940. His classroom contact withErnst and Berta in histology and neuroanatomy ledhim to spend the summer after his freshman yearworking in their laboratory. This scientific collabora-tion perservered throughout the remainder of Palay’smedical training and into his residency and was largelyresponsible for initiating his long and productive careeras a neuroanatomist.

In 1946 the Scharrers left Cleveland for the Depart-ment of Anatomy at the University of Colorado inDenver, where Ernst was appointed associate professor.During their nine-year stay in Denver, Berta, stillwithout a salaried academic appointment, was awardeda Guggenheim Fellowship and then a Special Fellow-ship of the NIH. This was an especially productiveperiod for Berta for she was able to complete her classicwork on the physiology of the insect neurosecretorysystem. The year 1955 was a milestone in Berta’sscientific career. The Scharrers left Denver to found theDepartment of Anatomy at the newly opening AlbertEinstein College of Medicine in the Bronx, New York,where Ernst had been appointed chairman. At the sametime the college appointed Berta full professor, award-ing her an academic appointment for the first time inher scientific career. Her previous lack of status reflectedthe prejudice against even the suggestion of nepotismthat prevailed in academic institutions of that era, aprejudice that, as mentioned above, strongly influencedErnst. But more importantly, it reflected the strongprejudice against women as serious scientists. AlthoughBerta’s appointment as professor under her husband’schairmanship was full-time, she received only half sal-ary. It was not until a decade later, when she was ap-pointed acting chair of the department following Ernst’saccidental death, that she began to receive full salary.

The Scharrers began their teaching duties almostimmediately upon arrival in the Bronx. Berta served ascourse director of histology. To assemble teaching mate-rial, recruit faculty, and design and administer thecourse de novo were demanding tasks that consumedher attention for several years, imposing a lull in herresearch activity. When she did return to her research,she initiated a series of studies of the structure of insectneurosecretory organs with the electron microscope,publishing her first paper on the ultrastructure of thecorpus cardiacum in 1963.

This phase of Berta’s research continued for the next15 years. During this period she observed that therough-surfaced endoplasmic reticulum and Golgi appa-ratus of insect neurosecretory cells produced membrane-bound secretory granules in the same manner as didconventional secretory cells, that the granules passeddown the axon via axonal transport, that the granuleswere discharged at the nerve endings in different ways,either by release into a neurochemal organ, whichenabled the neuropeptide to reach its target via the

hemolymph, or by release in close proximity of thetarget cells, a more direct pathway of communication.She also observed that complex structures with many ofthe characteristics of the vertebrate synapse werepresent along the surface of the axon of neurosecretorycells.

She had just begun these studies when she lost herhusband. In 1965 Ernst drowned while swimming inthe Atlantic on the Florida coast after the annualmeeting of the Anatomists held in Miami Beach. Bertaherself narrowly escaped the undertow that carriedErnst away. Declining to be considered for his succes-sor, she did assume the duties of acting chairperson.The next year Dominick Purpura was named to headthe department, and Berta could return to research andteaching. When Purpura left the department in 1976,Berta again undertook the duties of acting chairpersonuntil the following year when Peter Satir became thenew chairman.

In 1978, at age 72, Berta was appointed Distin-guished University Professor Emerita. Rather thanrest on her laurels, she continued her scientific careerfor the next 17 years until her death at 88 in 1995.

The pattern of Berta’s scientific output from 1974until 1995 informs us about her impressive qualities asa scientist. Starting from 1974 her output of originalresearch papers was interrupted, indicating that shehad set aside her own research program, and she beganto devote her time to writing scientific reviews. In the35 years prior to 1975, she had published a total of onlyfive reviews. During the six-year period from 1975 until1981 she published ten reviews, and it looked as thoughthis activity would lay claim to her scientific talent fromthat point on, particularly since she was a gifted sciencewriter. However, 1982 marked a surprising turningpoint in her career; she began a new series of researchprojects in collaboration with the Danish scientists,Georg and Bente Hansen, and with George Stefano.These she continued until the time of her death. At age76, she had reactivated her career as a scientificinvestigator. In the next eight years she publishedseven scientific articles on the identification and func-tion of neuropeptides in invertebrates. She and herco-workers showed that the same neuropeptides identi-fied in vertebrates were also present in invertebrateswhere they functioned similarly. She rightfully con-cluded that the commonality, between the vertebrateand invertebrate phyla, of the enzyme systems for thesynthesis and metabolic breakdown of neuropeptidesand for the receptor mechanisms to implement theirmode of action was strong evidence of the long evolution-ary history of the neurosecretory system. Her grasp ofthe fundamentals and implications of the subject ofneuropeptides was so comprehensive that she was ableto write five critical reviews about this subject duringthe same eight-year period.

Incredible though it may seem, in 1990 at age 84,Berta moved to the forefront of yet another newlyemerging field: neuroimmunology. In the remaining sixyears of her life, despite being plagued by increasingphysical disability, she pursued her interest in this areaof research with her customary vigor, adding 11 re-search papers and three reviews to her bibliography aswell as serving as associate editor of Advances inNeuroimmunology. She and her colleagues showed that

2 S.L. WISSIG

the neuropeptides and receptors they had previouslystudied play an additional important role in regulationof the immune response in invertebrate species as theydo in vertebrates.

On July 23, 1995 illness finally brought an end toBerta’s illustrious scientific career.

Berta gained worldwide recognition for her scientificachievements. She was named honorary member ofeight national and foreign scientific societies. She re-ceived honorary degrees from the Universities of Gies-sen, Frankfurt, and Salzburg in Europe, the Universityof Calgary in Canada, and from a number of Americaninstitutions including the University of North Carolina,the State University of New York (Old Westbury),Smith and Mount Holyoke Colleges, and Northwestern,Harvard, and Yeshiva Universities. Major awards shereceived included the Kraepelin Gold Medal for herstudies in neuroendocrinology, the F.C. Koch Award ofthe Endocrine Society, the Henry Gray and CentennialAwards of the American Association of Anatomists, theSchleiden Award of the Deutsche Akademie fur Natur-forscher Leopoldina, and the Order of Merit of the FreeState of Bavaria among others. She served as Presidentof the American Association of Anatomists in 1979–80.She was elected to membership in the American Acad-emy of Arts and Sciences and the National Academy ofSciences. President Reagan awarded her the NationalMedal of Science in 1983.

Both she and Ernst showed great concern for theirstudents and colleagues. As teachers they inspired anumber of their medical students to undertake careersin research. Palay has already been mentioned; theneuroanatomist, Harvey Karten, and the endocrinolo-gist, Jesse Roth, are other examples. With these indi-viduals, associations that began in the classroom ma-tured into lifelong friendships.

Some of Berta’s most meaningful work was done inediting for the Proceedings of the National Academy ofSciences, Advances in Neuroimmunology, and Cell andTissue Research (formerly Zeitschrift fur Zellforsch-ung). Scientific quality and integrity, blended withconcern for authors, characterized her editing as sheworked tirelessly, during health and illness, not only tojudge their work but also to assist and guide them. Shesucceeded as an editor because she did not let her ownscientific achievements efface her humility. On occa-sion, her humility could lead to an amusing incident.When she was at Harvard to attend the ceremony atwhich she was to receive her honorary degree, theparticipants in academic regalia were assembling forthe procession. Harvard had cautioned Berta prior tothe ceremony that recipients of the degree were not todivulge that they had been selected before the awardwas made. The Dean of Albert Einstein medical school,a division of Yeshiva University, was in the audienceattending the ceremony. Spotting Berta, her dean cameup to her and asked what she was doing there. Mindfulof Harvard’s request, all she could do was shrug hershoulders noncommittally. It is a matter of record thatYeshiva University awarded her an honorary degreethe following year. This same humility and concern forothers kept her working to the best of her ability until afew days before her death.

A number of Berta’s achievements can be singled outas highlights of her distinguished career. To begin with,

she was fortunate to have had the opportunity and thedetermination to work at an undiminished creativepace from the completion of her education until hereighty-ninth year, a period of 65 years. Then fate andmatrimony positioned her at the birth of a very signifi-cant branch of biological science: the crosscommunica-tion between the nervous and endocrine systems. Whilestill a neophyte in science, she and her husband had thevision to imagine where their anatomical experimentsmight lead. Once the authenticity of their insight intothe significance of neurosecretion was established, sherode the crest of the wave of the research that docu-mented its physiologic workings, primarily in inverte-brates. In order to keep pace with ensuing develop-ments in links between the nervous system and theother major integrating systems of the body, i.e., thehormone and immune systems, she moved from neurse-cretion successively into neuropeptides and neuroimmu-nology. Remarkably, age seemed no barrier to herentrance into newly emerging research areas. Finally,and probably of singular importance to her colleaguesin the American Association of Anatomists, is the factthat her experimental findings were chiefly based onmicroscopic structure, i.e., she succeeded in sciencebasically as a morphologist. Thus, for her notableaccomplishments, she has earned a place of honoramong anatomists.

SELECTED SCIENTIFIC PUBLICATIONSOF BERTA SCHARRER

Original Articles*Early Work in Germany(as B. Vogel) 1931 Uber die Beziehungen zwischen Sussgeschmack

und Nahrwert von Zuckern und Zuckeralkoholen bei der Honig-biene. Z. vergleichende Physiol., 14:273–347.

(as B. Vogel) 1931 Uber die Beziehungen zwischen Sussgeschmackund Nahrwert von Zuckern bei der Honigbiene. Sitzungsber. Ges.Morph. Physiol., 40:63–65.

(as B. Vogel) 1933 Vergleichend-morphologische Untersuchungen anHuhner-und Recurrensspirochaten. Sitzungsber. Ges. Morph.Physiol. Munchen., 42:1–5.

1934 Uber die Verweildauer von Huhnerspirochaten im Zentralnerven-system von Huhnern und Tauben. Z. Hygiene, 116:206–208.

1934 Ein serumfreier Nahrboden fur Huhnerspirochaten. Zentrabl.Bakteriol. I. Orig., 132:243–244.

(with F. Jahnel) 1935 Ein Beitrag zur Frage der Syphilisempfindlich-keit im Tierreich, insbesondere verschiedener Mausearten. Derma-tol. Z., 71:1–6.

1935 Uber das Hanstromsche Organ X bei Opisthobranchiern. Pubbl.Stn. Zool. Napoli., 15:132–142.

1935 Uber das Verhalten von Tauben gegenuber Infektionen mitverschiedenen Huhnerspirochatenstammen. Z. Hygiene, 117:163–170.

1935 Uber Spirochaeta (Treponema) minutum Dobell bei amphibien.Zool. Anzeiger, 111:1–7.

1935 Uber die Feststellung von Dickenunterschieden an lebendenmikroskopischen objekten, dargelegt am Beispiel der Unterschei-dung von Huhner- und Recurrensspirochaten. Arch. Protisten-kunde, 85:87–99.

1936 Uber ‘‘Drusen-Nervenzellen’’ im Gehirn von Nereis virens Sars.Zool. Anzeiger, 113:299–302.

Establishment of the Basis of Invertebrate Neurosecretion1937 Uber sekretorisch tatige Nervenzellen bei wirbellosen Tieren.

Naturwissenschaften, 25:131–138.

*Important original articles are cited in full in this section. Selectedreviews and books are cited separately below. With some exceptions,book reviews, meeting reviews, etc., authored or co-authored by BertaScharrer, are omitted. Articles about her are cited at the end of thelisting.

3A TRIBUTE TO BERTA SCHARRER

(with E. Hadorn) 1938 The structure of the ring-gland (corpusallatum) in normal and lethal larvae of Drosophila melanogaster.Proc. Natl. Acad. Sci. U.S.A., 24:236–242.

1939 The differentiation between neuroglia and connective tissuesheath in the cockroach (Periplaneta americana). J. Comp. Neu-rol., 70:77–88.

1941 Neurosecretion. II. Neurosecretory cells in the central nervoussystem of cockroaches. J. Comp. Neurol., 74:93–108.

1941 Neurosecretion. III. The cerebral organ of the nemerteans. J.Comp. Neurol., 74:109–130.

1941 Neurosecretion. IV. Localization of neurosecretory cells in thecentral nervous system of Limulus. Biol. Bull., 81:96–104.

(with E. Scharrer) 1944 Neurosecretion VI. A comparison between theintercerebralis-cardiacum-allatum system of the insects and thehypothalamo-hypophyseal system of the vertebrates. Biol. Bull.,87:242–251.

1945 Experimental tumors after nerve section in an insect. Proc. Soc.Exp. Biol. Med., 60:184–189.

1946 The role of the corpora allata in the development of Leucophaeamaderae (Orthoptera). Endocrinology, 38:35–45.

1946 The relationship between corpora allata and reproductive organsin adult Leucophaea maderae (Orthoptera). Endocrinology, 38:46–55.

1948 The prothoracic glands of Leucophaea maderae (Orthoptera).Biol. Bull. 95:186–198.

1949 Gastric cancer experimentally induced in insects by nerveseverance. J. Natl. Cancer Inst., 10:375–376.

1951 Tumors in invertebrates. In: Year Book of Pathology, Vol. 1950. pp57–61. Chicago, IL: The Year Book Medical Publishers.

1951 The woodroach. Sci. Am., 185:58–62.1952 Neurosecretion. XI. The effects of nerve section on the intercere-

bralis-cardiacum-allatum system of the insect Leucophaea mad-erae. Biol. Bull., 102:261–272.

1953 Insect tumors induced by nerve severance: Incidence and mortal-ity. Cancer Res., 13:73–76.

1953 Metabolism and mortality in insects with gastrointestinal tu-mors induced by nerve severance. J. Natl. Cancer Inst., 13:951–954.

(with E. Scharrer) 1953 Symposium on Neurosecretion at Naples,Italy, May 11–16. Science, 118:579–580.

1954 Neurosecretion in invertebrates: A survey. Publ. Stn. Zool.Napoli, 24(Suppl):38–40.

1955 Castration cells in the central nervous system of an insect(Leucophaea maderae, Blattaria). Trans. N.Y. Acad. Sci. Ser. II,17:520–525.

1956 Correlations endocrines dans la reproduction des insectes. Ann.Sci. Natl., Zool. ser. II, vol. 18: 231–234.

(with M. von Harnack) 1958 Histophysiological studies on the corpusallatum of Leucophaea maderae. I. Normal life cycle in male andfemale adults. Biol. Bull., 115:508–520.

Ultrastructural Studies on InvertebrateNeurosecretory SystemsNeurosecretion. XIII. 1963 The ultrastructure of the corpus cardiacum

of the insect Leucophaea maderae. Z. Zellforsch., 60:761–796.1964 The ultrastructure of the corpus allatum of Blaberus craniifer

(Blattaria). Am. Zool., 4:327–328.1964 Histophysiological studies on the corpus allatum of Leucophaea

maderae. IV. Ultrastructure during normal activity cycle. Z.Zellforsch., 62:125–148.

1964 The fine structure of blattarian prothoracic glands. Z. Zellforsch.,64:301–326.

1965 The fine structure of an unusual hemocyte in the insect Grompha-dorhina portentosa. Life Sci., 4:1741–1744.

1966 Ultrastructural study of the regressing prothoracic glands ofblattarian insects. Z. Zellforsch., 69:1–21.

(with E. Harper and S. Seifter) 1967 Electron microscopic andbiochemical characterization of collagen in blattarian insects. J.Cell Biol., 33:385–393.

1967 Ultrastructural specializations of neurosecretory terminals inthe corpus cardiacum of cockroaches. Am. Zool., 7:721–722.

1968 Neurosecretion. XIV. Ultrastructural study of sites of release ofneurosecretory material in blattarian insects. Z. Zellforsch., 89:1–16.

(with S.B. Kater) 1969 Neurosecretion. XV. An electron microscopicstudy of the corpora cardiaca of Periplaneta americana afterexperimentally induced hormone release. Z. Zellforsch. 95:177–186.

(with S. Wurzelmann) 1969 Ultrastructural study on nuclear-cytoplasmic relationships in oocytes of the African lungfish,

Protopterus aethiopicus. I. Nucleolo-cytoplasmic pathways. Z.Zellforsch., 96:325–343.

(with S. Wurzelmann) 1969 Ultrastructural study on nuclear-cytoplasmic relationships in oocytes of the African lungfish,Protopterus aethiopicus. II. The microtubular apparatus of thenuclear envelope. Z. Zellforsch., 101:1–12.

1970 Ultrastructural study of the sites of origin and release of acellular product in the corpus allatum of insects. Proc. Natl. Acad.Sci., 66:244–245.

1971 Histophysiological studies on the corpus allatum of Leucophaeamaderae. V. Ultrastructure of sites of origin and release of adistinctive cellular product. Z. Zellforsch., 120:1–16.

1972 Cytophysiological features of hemocytes in cockroaches. Z.Zellforsch., 129:301–319.

(with S. Wurzelman) 1974 Observations on synaptoid vesicles in insectneurons. Zool. Jahrbucher Abt. Physiol., 78:387–396.

(with S. Wurzelmann) 1977 Neurosecretion. XVI. Protrusions ofbounding membranes of neurosecretory granules. Cell TissueRes., 184:79–85.

Neuropeptides and Neuroimmunology(with G.B. Stefano and P. Assanah) 1982 Opioid binding sites in the

midgut of the insect Leucophaea maderae (Blattaria). Life Sci.,31:1397–1400.

(with G.N. Hansen and B.L. Hansen) 1987 Gastrin/CCK-like immuno-reactivity in the corpus cardiacum-corpus allatum complex of thecockroach Leucophaea maderae. Cell Tissue Res., 248:595–598.

(with G.N. Hansen and B.L. Hansen) 1988 Diversity of prolactinsystems in the insect Leucophaea maderae: Use of antiserumpolyclonality for immunocytochemical detection of neuropeptideheterogeneity. Cell Tissue Res., 252:557–563.

(with G.B. Stefano and M.K. Leung) 1988 Opioid mechanisms ininsects with special attention to Leucophaea maderae. Cell Mol.Neurobiol., 8:269–284.

(with G.B. Stefano, M.K. Leung, et al.) 1989 Evidence for the involve-ment of opioid neuropeptides in the adherence and migration ofimmunocompetent invertebrate hemocytes. Proc. Natl. Acad. Sci.U.S.A., 86:6307–6311.

(with G.B. Stefano and P. Cadet) 1989 Stimulatory effects of opioidneuropeptides on locomotory activity and conformational changesin invertebrate and human immunocytes: Evidence for a subtypeof S receptor. Proc. Natl. Acad. Sci. U.S.A., 86:6307–6311.

(with G.N. Hansen, B.L. Hansen, et al.) 1990 Immunocytochemicallocalization and immunochemical characterization of an insulin-related peptide in the insect Leucophaea maderae. Cell TissueRes., 259:265–273.

(with T.K. Hughes, Jr., E.M. Smith, et al.) 1990 Interaction ofimmunoactive monokines (interleukin 1 and tumor necrosis fac-tor) in the bivalve mollusc Mytilus edulis. Proc. Natl. Acad. Sci.U.S.A., 87:4426–4429.

(with M.A. Shipp, G.B. Stefano, et al.) 1990 Downregulation ofenkephalin-mediated inflammatory responses by CD10/neutralendopeptidase 24.11. Nature, 347:394–396.

(with G.B. Stefano, P. Cadet, et al.) 1990 A neuroimmunoregulatory-like mechanism responding to stress in the marine bivalveMytilus edulis. Brain, Behav. Immun., 4:323–329.

(with G.B. Stefano, P. Cadet, et al.) 1990 Neural activation of thecellular immune system involving an opioid mechanism in themollusc Mytilus edulis. Ann. N.Y. Acad. Sci., 594:494–498.

(with G.B. Stefano and M.A. Shipp) 1991 A possible immunoregulatoryfunction for (Met)-enkephalin-Arg6-Phe7 involving human andinvertebrate granulocytes. J. Neuroimmunol., 31:97–103.

(with M.A. Shipp, G.B. Stefano, et al.) 1991 CD10 (CALLA, commonacute lymphoblastic leukemia antigen)/neutral endopeptidase24.11 (NEP, ‘‘enkephalinase’’): Molecular structure and role inregulating met-enkephalin mediated inflammatory responses.Adv. Neuroimmunol., 1:139–149.

(with G.B. Stefano, P. Melchiorri, et al.) 1992 (D-Ala2) deltorphin Ibinding and pharmacological evidence for a special subtype ofdelta opioid receptor on human and invertebrate immune cells.Proc. Natl. Acad. Sci. U.S.A., 89:9316–9320.

(with G.B. Stefano, A. Digenis, et al.) 1993 Opiatelike substances in aninvertebrate: A novel opiate receptor on invertebrate and humanimmunocytes, and a role in immunosuppression. Proc. Natl. Acad.Sci. U.S.A., 90:11099–11103.

(with G.B. Stefano, M.K. Leung, et al.) 1993 Autoimmunoregulationand the importance of opioid peptides. Ann. N.Y. Acad. Sci.712:92–101.

(with D. Sonetti, E. Ottaviani, et al.) 1994 Microglia in invertebrateganglia. Proc. Natl. Acad. Sci. U.S.A., 91:9180–9184.

4 S.L. WISSIG

(with G.B. Stefano) 1995 The presence of the µ3 opiate receptor ininvertebrate neural tissue. Comp. Biochem. Physiol., Part C,Pharmacol. Toxicol. Endocrinol. 113:369–373.

(with G.B. Stefano, M.K. Leung, et al.) 1995 Effect of prolongedexposure to morphine on responsiveness of human and inverte-brate immunocytes to stimulating molecules. J. Neuroimmunol.63:175–181.

(with L. Paemen, E.M. Smith, et al.) 1996 The presence and effects ofmammalian signal molecules in immunocytes of the insect Leuco-phaea moderae. Cell Tissue Res. 283:93–97.

Selected Reviews and Commentaries(with E. Scharrer) 1937 Uber Drusen-Nervenzellen und neurosekreto-

rische Organe bei Wirbellosen und Wirbeltieren. Biol. Rev., 12:185–216.

(with E. Scharrer) 1941 Endocrines in invertebrates. Physiol. Rev.,21:383–409.

(with E. Scharrer) 1945 Neurosecretion., Physiol. Rev., 25:171–181.1948 Hormones in insects. In: The Hormones, Physiology, Chemistry,

and Applications, Vol. I. G. Pincus, K.V. Thimann, eds. New York:Academic Press, pp. 121–158.

1952 Hormones in insects. In: The Action of Hormones in Plants andInvertebrates. K.V. Thimann, ed. New York: Academic Press,1952, pp. 125–169.

1953 Comparative physiology of invertebrate endocrines. Ann. Rev.Physiol., 15:457–472.

1965 Recent progress in the study of neuroendocrine mechanisms ininsects. Arch. Anat. Microsc. Morphol Exp., 54:331–342.

1967 The neurosecretory neuron in neuroendocrine regulatory mecha-nisms. Am. Zool., 7:161–169.

1968 Neuroendocrine factors in the control of reproduction. In: Repro-duction and Sexual Behavior. M. Diamond, ed. Bloomington, IN:Indiana Univ. Press, pp. 145–149.

1970 General principles of neuroendocrine communication. In: TheNeurosciences: Second Study Program. F.O. Schmitt, ed. NewYork: Rockefeller Univ. Press, pp. 519–529.

(with M. Weitzman) 1970 Current problems in invertebrate neurose-cretion. Proc. Vth Int. Symp. Neurosecretion, Berlin.

1975 The role of neurons in endocrine regulation: A comparativeoverview. Am. Zool., 15:(Suppl. 1)7–11.

1975 The concept of neurosecretion and its place in neurobiology. In:The Neurosciences: Paths of Discovery. F.G. Worden, J.P. Swaxey,et al., ed. Cambridge, MA. The MIT Press, 1975, pp. 231–243.

1975 Neurosecretion and its role in neuroendocrine regulation. In:Pioneers in Neuroendocrinology. J. Meites, B.T. Donovan, et al.,eds. New York and London: Plenum Press, pp. 257–265.

1976 Neurosecretion: Comparative and evolutionary aspects, In:Progress in Brain Research. M.A. Corner and D.F. Swash, eds.Amsterdam: Elsevier Publishing Co., pp. 125–137.

1977 Evolutionary aspects of neuroendocrine control processes. In:Reproductive Behavior and Evolution. J.S. Rosenblatt and B.R.Komisaruk, eds. New York and London: Plenum Press, pp.111–124.

1977 Peptides in neurobiology: Historical introduction. In: Peptides inNeurobiology. H. Gainer, ed. New York and London: PlenumPress, pp. 1–8.

1978 An evolutionary interpretation of the phenomenon of neurosecre-tion. Forty-Seventh James Arthur Lecture on the Evolution of theHuman Brain, 1–17. New York: The American Museum of NaturalHistory, 1978.

1978 Peptidergic neurons: Facts and trends. Gen. Comp. Endocrinol.,34:50–62.

1979 Neurosecretion and neuroendocrinology in historical perspec-tive. In: Hormonal Proteins and Peptides. C.H. Li, ed. 7:279–292.

1981 Recent results on the neuroendocrine system of Leucophaea. In:Current Topics in Insect Endocrinology and Nutrition. G.Bhaskaran, S. Friedman, et al., eds. New York, London: PlenumPress, pp. 47–52.

1981 Peptidergic Neurons: Retrospect and Prospect. Verh. Anat. Ges.,75:977–978.

1987 Insects as model in neurendocrine research. Ann. Rev. Entomol.,32:1–16.

1987 Neurosecretion: Beginnings and new directions in neuropeptideresearch. Ann. Rev. Neurosci., 10:1–17.

(with G.N. Hansen and B.L. Hansen) 1988 Diversity of prolactinsystems in the insect Leucophaea maderae: Use of antiserumpolyclonality for immunocytochemical detection of neuropeptideheterogeneity. Cell Tissue Res., 252:557–563.

(with G.B. Stefano and M.K. Leung) 1988 Opioid mechanisms ininsects with special attention to Leucophaea maderae. Cell Mol.Neurobiol., 8:269–284.

Neurobiology of opioids in Leucophaea maderae. In: Insect Models inResearch. I. Huber, ed. Boca Raton, FL: CRC Press, pp. 85–102.

1990 The neuropeptide saga. Am. Zool., 30:887–895.1991 Neuroimmunology: The importance and role of a comparative

approach. Adv. Neuroimmunol., 1:1–6.1992 Recent progress in comparative neuroimmunology. Zool. Sci.

(Japan), 9:1097–1100.(with G.B. Stefano) 1994 Endogenous morphine and related opiates, a

new class of chemical messengers. Adv. Neuroimmunol. 4:57–67.(with G.B. Stefano, T.O. Bilfiger, et al.) 1996 A novel view of opiate

tolerance. Adv. Neuroimmunol. 6:265–277.

Books(with E. Scharrer) 1963 Neuroendocrinology. New York, London:

Columbia Univ. Press, 289 p. (The seminal book by Ernst andBerta Scharrer, establishing the field of Neuroendocrinology.)

Aspects of Neuroendocrinology (W. Bargman and B. Scharrer, eds.)1970 Proc. Vth Int. Symp. Neurosecretion. Berlin-Heidelberg:Springer-Verlag, 380 p.

Neurosecretion and Neuroendocrine Activity (W. Bargman, A. Oksche,A. Polenov, B. Scharrer, eds.) 1978 Berlin-Heidelberg: SpringerVerlag, 411 p.

Neuropeptides and Immunoregulation. B. Scharrer et al., eds. 1994Berlin-Heidelberg: Springer-Verlag, 174 p.

Other Works1987 Pioneers in Modern Biology, Berta Scharrer: A Partner in the

Discovery of Neurohormones. VHS Videotape. K.R. Porter Endow-ment in Cell Biology. Philadelphia, PA: Univ. of Pennsylvania.(Berta Scharrer and others discuss the origins of the concept ofneurosecretion.)

1987 Against The Tide. Women in Cell Biology. VHS Videotape. K.R.Porter Endowment in Cell Biology. Philadelphia, PA: Universityof Pennsylvania. (In the first part of this videotape, BertaScharrer discusses her early experiences as a woman in science.)

Works about Berta Vogel Scharrer1965 Neuroendocrine Mechanisms. Farrell, G., ed. Science 147:1136.1978 Neuroendocrinology: Pioneering efforts. (letter). G.W. Szilard, ed.

Science 200:1107 (June 9 issue).1980 Citation for the Fred Conrad Koch Award of the Endocrine

Society to Berta Scharrer. Endocrinology 107:364–365. (An excel-lent brief review of Berta Scharrer’s scientific accomplishments to1980.)

1982 Henry Gray Award in Anatomical News, Series 3, No. 8, May,1982 (Amer. Assoc. Anatomists, Publishers). (A brief review ofBerta Scharrer’s career.)

1989 On Journeys Well Traveled. In: Einstein. Millen, S.K., ed. Bronx,NY: Office of Public Affairs, Albert Einstein College of Medicine,pp. 3–6. (An overview of Berta Scharrer’s life and work withquotations from an interview with Berta Scharrer.)

Martin, D. 1995 Roach Queen Retires. N.Y. Times (Feb. 9, 1995),B1–B2.

Saxon, Wolfgang. 1995 Berta Scharrer, 88, research scientist androach expert. N.Y. Times (July 25, 1995), A13.

Kreeger, Karen Y. 1995 Pioneering neuroscientist Berta Vogel Schar-rer dies. Scientist (Sept. 4, 1995), 17.

Siebert, Charles. 1995 What the roaches told her. N.Y. Times Maga-zine (December 31, 1995), 26–27.

5A TRIBUTE TO BERTA SCHARRER