University of Southern Denmark Hans Bluntschli in Berne Researching reproduction in hedgehog tenrecs (Afrosoricida, Tenrecidae) Carter, Anthony M Published in: Journal of Morphology DOI: 10.1002/jmor.20988 Publication date: 2019 Document version: Accepted manuscript Citation for pulished version (APA): Carter, A. M. (2019). Hans Bluntschli in Berne: Researching reproduction in hedgehog tenrecs (Afrosoricida, Tenrecidae). Journal of Morphology, 280(6), 841-848. https://doi.org/10.1002/jmor.20988 Go to publication entry in University of Southern Denmark's Research Portal Terms of use This work is brought to you by the University of Southern Denmark. Unless otherwise specified it has been shared according to the terms for self-archiving. If no other license is stated, these terms apply: • You may download this work for personal use only. • You may not further distribute the material or use it for any profit-making activity or commercial gain • You may freely distribute the URL identifying this open access version If you believe that this document breaches copyright please contact us providing details and we will investigate your claim. Please direct all enquiries to [email protected] Download date: 04. Oct. 2021 Carter Anthony (Orcid ID: 0000-0002-8359-2085) Hans Bluntschli in Berne: researching reproduction in hedgehog tenrecs (Afrosoricida, Tenrecidae) Anthony M. Carter Cardiovascular and Renal Research, Institute of Molecular Medicine, University of Southern Denmark, J. B. Winsloews Vej 21, DK-5000 Odense, Denmark Correspondence Anthony M. Carter, Cardiovascular and Renal Research, Institute of Molecular Medicine, University of Southern Denmark, J. B. Winsloews Vej 21, DK-5000 Odense, Denmark. Email: [email protected] Research Highlights Tenrec reproduction was the main aspect of Bluntschli’s research after was obliged to leave Frankfurt for Berne His group described unique features that included non-antral follicles, intraovarian fertilization and absence of a morula stage Abstract The Swiss anatomist Hans Bluntschli is best known as a primatologist. Yet his focus during his later years in Berne was on reproduction in Malagasy tenrecs. This research was done with two graduate students, Robert Goetz and Fritz Strauss; all three had been obliged to leave Germany after the National Socialists came to power. Unique features of reproduction in tenrecs included non-antral follicles, intrafollicular fertilization, eversion of the corpus luteum and polyovulation. The fertilized egg formed a blastula that developed into a blastocyst; there was no morula stage. A false placental cushion developed in the endometrium opposite the implantation site. Placentation was complex and included development of a prominent haemophagous organ. These findings are discussed in relation to current concepts of mammalian phylogeny that place tenrecs and golden moles in the same order and as close relatives to elephant shrews and the aardvark. KEYWORDS blastula, history of science, non-antral follicles, placentation, polyovulation This is the author manuscript accepted for publication and has undergone full peer review but has not been through the copyediting, typesetting, pagination and proofreading process, which may lead to differences between this version and the Version of Record. Please cite this article as doi: 10.1002/jmor.20988 This article is protected by copyright. All rights reserved. 1 INTRODUCTION In 1931, Hans Bluntschli, renowned primatologist and Director of the prestigious Senckenberg Research Institute, embarked on an expedition to Madagascar (Fig. 1). Two years later, and shortly after his return, he was relieved of his duties in Frankfurt by the National Socialist regime. He then was called to the chair of anatomy at Berne. Although he had collected primate material on Madagascar, his first priority was to describe the reproductive biology of tenrecs. To this end, he supervised research by Robert Goetz, who had been denied a licence to practise medicine in Germany, and Fritz Strauss, who had fled to Switzerland because of his Jewish parentage. Together they described some remarkable features of reproduction in tenrecs such as the absence of antral follicles, lack of a morula stage, large litter size and unique placentation. This work occupied Bluntschli during his most productive years at Berne, yet has been overlooked by previous biographers (Cener, 1990; Greif & Schmutz, 1995). Tenrec biology attracted scant attention until recently, when molecular phylogenetics shook the mammalian tree (Madsen et al., 2001; Murphy, Eizirik, Johnson, et al., 2001). It focussed interest on a superorder of mammals that included tenrecs, placing them near to the root of the eutherian tree (Carter, 2001). All three species studied by Bluntschli, Goetz and Strauss were from the same subfamily of tenrecs (Tenrecinae) for which they used the synonym Centetinae (McKenna & Bell, 1997). Tenrecs were seen by them to be very primitive insectivores as were golden moles (Bluntschli, 1938). Based on molecular phylogenetics, tenrecs and golden moles are now grouped in the order Afrosoricida (Madsen et al., 2001). In contrast to what was known in Bluntschli’s time, they are thought to belong to a different lineage than other insectivores, such as hedgehogs, shrews and moles. Today, the three subfamilies of Malagasy tenrecs and the African otter shrews comprise the suborder Tenrecomorpha (Everson, Soarimalala, Goodman, & Olson, 2016). This review has a dual purpose. The first is to recapitulate the careers of three scientists and their intersection in the shadow of the Third Reich (Hildebrandt & Redies, 2012). The other is to review the unique aspects of tenrec biology they described in the late 1930’s and 1940’s and, where relevant, update them with recent data. This story is a timely reminder that research does not operate in a vacuum but must be viewed in the context of contemporary politics and societal norms (Hildebrandt & Redies, 2012). 2 THE SCIENTISTS 2 This article is protected by copyright. All rights reserved. The principal sources for Bluntschli are an obituary (Strauss, 1964), more recent biographies dealing with him as anatomist (Cener, 1990) and morphologist (Greif & Schmutz, 1995) and detailed accounts of his dismissal from Frankfurt under the National Socialist regime (Benzenhöfer, 2014; Benzenhöfer & Hack- Molitor, 2012). The most comprehensive account of Goetz deals principally with his career after leaving Berne, but includes his own thoughts on the time spent there (Konstantinov, 2000). There is a useful obituary of Strauss (Weber, 1995). Hermann Georg Hans Bluntschli was born at Frankfurt on Main on 19 February 1877. His father was a renowned Swiss architect. Bluntschli studied medicine at the University of Zurich but developed symptoms of a hereditary hearing impediment (otosclerosis) that he knew would hinder a clinical career. In consequence, after his final exams, he moved to the Anatomical Institute at Heidelberg. There, under the guidance of Max Fürbringer (1846-1920), a student of Carl Gegenbauer (1826-1903), he wrote a dissertation on the liver of the Queensland lungfish Neoceratodus forsteri (Bluntschli, 1904). He then returned to Zurich with a view to studying the comparative anatomy of primates. His thesis (Habilitation) was about the branches of the femoral artery in catarrhine primates (Bluntschli, 1906). In 1912 he made an expedition to Brazil to collect material from Neotropical primates, following in the footsteps of the eminent Swiss-Brazilian zoologist Emil August Goeldi (1859-1917). On his return, he was recruited by the Senckenberg Research Institute in Frankfurt, becoming a full professor in 1914. Here he did his most important primatological work (Greif & Schmutz, 1995). From 1931 to 1932, he embarked on an expedition to Madagascar, where he collected lemurs and tenrecs with the intent of studying their embryology. Not long after his return, he was relieved of his duties by the National Socialist regime (Benzenhöfer & Hack- Molitor, 2012). He was, however, called to the chair of anatomy in Berne, where he remained until his retirement in 1947. Bluntschli favoured a holistic approach to anatomy (Benzenhöfer & Hack-Molitor, 2012) and in 1938 founded a new journal Bio-Morphosis, with the aim of fostering the exchange of ideas between morphologists and physiologists (Strauss, 1964). A considerable part of the tenrec work was published there. Hans Bluntschli died at Berne on 13 July 1962. Robert Hans Goetz was born in Frankfurt 17 April 1910. He read medicine at the University of Frankfurt and while still a student invented a finger plethysmograph (Goetz, 1933). In 1934, after completing his finals, he was declared “politically unreliable” and denied a medical license. Meanwhile, Goetz had married Verena Bluntschli and his father-in-law gave him the opportunity to work on the tenrec material in Berne 3 This article is protected by copyright. All rights reserved. (Konstantinov, 2000). In 1937 he went to South Africa, where he had to retake his medical exams before practising medicine. During the Cape Town years, he conducted innovative field studies on the cardiovascular physiology of the giraffe (Mitchell, 2008). In 1957 Goetz, by then a cardiac surgeon, settled in the United States. While at the Albert Einstein College of Medicine, New York, he performed the first successful coronary bypass operation (Konstantinov, 2000). Throughout his career, Goetz was supported by his wife, who was a physician trained in Geneva