CJ Van Der Horst on Pregnancy and Menstruation in Elephant Shrews
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University of Southern Denmark Classics revisited: C. J. van der Horst on pregnancy and menstruation in elephant shrews Carter, A M Published in: Placenta DOI: 10.1016/j.placenta.2018.05.010 Publication date: 2018 Document version: Accepted manuscript Document license: CC BY-NC-ND Citation for pulished version (APA): Carter, A. M. (2018). Classics revisited: C. J. van der Horst on pregnancy and menstruation in elephant shrews. Placenta, 67, 24-30. https://doi.org/10.1016/j.placenta.2018.05.010 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 Accepted Manuscript Classics revisited: C. J. van der Horst on pregnancy and menstruation in elephant shrews A.M. Carter PII: S0143-4004(18)30082-1 DOI: 10.1016/j.placenta.2018.05.010 Reference: YPLAC 3828 To appear in: Placenta Received Date: 23 March 2018 Revised Date: 14 May 2018 Accepted Date: 25 May 2018 Please cite this article as: Carter AM, Classics revisited: C. J. van der Horst on pregnancy and menstruation in elephant shrews, Placenta (2018), doi: 10.1016/j.placenta.2018.05.010. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. ACCEPTED MANUSCRIPT 1 Classics Revisited: C. J. van der Horst on pregnancy and menstruation in elephant shrews 2 A. M. Carter * 3 Cardiovascular and Renal Research, Institute of Molecular Medicine, University of Southern Denmark, 4 Odense 5 *Corresponding author: Anthony M. Carter. Tel: +45 6550 3716; Email: [email protected] 6 Running title: Menstruation in elephant shrews 7 No. words (abstract): 161 (1020 characters with spaces) 8 No. words (total): 3869 (24,967 characters with spaces) 9 Figures 4 10 Tables 1 MANUSCRIPT ACCEPTED 1 ACCEPTED MANUSCRIPT 11 Abstract 12 Background: Menstruation occurs only in higher primates, some bats, the spiny mouse and the elephant 13 shrew. Our knowledge of the latter species is due to work by C. J. van der Horst. 14 Findings: Changes in the uterine stroma are initially similar in fertile and infertile cycles and are confined to 15 a small area. In pregnant animals, the presence of the conceptus causes further development to an 16 implantation chamber. In infertile cycles an outgrowth of highly glandular stroma (a polyp) appears. With 17 decline of the corpora lutea it is shed in a process equivalent to menstruation. Van der Horst described the 18 further development of the placenta and a decidua pseudocapsularis in pregnant animals. In addition he 19 built a unique collection that has thrown light on embryonic development and placentation in other South 20 African mammals. 21 Conclusions: The changes in endometrial stromal cells during the menstrual cycle appear similar between 22 primates and the elephant shrew and deserve to be studied at the molecular level. MANUSCRIPT 23 Key words: Afrotheria; History of Science; Macroscelidea; Menstrual cycle; Placentation; Polyovulation ACCEPTED 2 ACCEPTED MANUSCRIPT 24 Highlights 25 • C. J. van der Horst was a distinguished marine biologist and an authority on acorn worms 26 • In later life he turned to the embryology of South African mammals 27 • An important contribution was his description of menstruation in elephant shrews 28 • His collection was used to describe early development and placentation in other South African 29 mammals 30 • Menstruation in elephant shrews is discussed in the context of recent work on endometrial stromal 31 cells MANUSCRIPT ACCEPTED 3 ACCEPTED MANUSCRIPT 32 1. Introduction 33 Human reproduction has many unique features [1]. One is menstruation, the shedding of decidua at the 34 end of an infertile cycle with accompanying loss of blood. Decidualization involves a change in the size, 35 shape and properties of the connective tissue cells of the endometrium (stromal fibroblasts) in preparation 36 for embryo implantation [2]. In most mammals, decidualization does not occur until there is an embryonic 37 signal. But in humans decidualization is triggered by a maternal signal in the second half of the menstrual 38 cycle. The decidua will be useful if there is a pregnancy, but otherwise is shed. 39 Menstruation is known to occur in haplorrhine primates (Old World monkeys and apes) and some 40 Neotropical primates. It has also been documented in phyllostomid and molossid bats [3] and the Cairo 41 spiny mouse ( Acomys cahirhinus ) [4]. However, the best-known example of menstruation in non-primates 42 concerns a distantly related order, the elephant shrews or sengis (Macroscelidea). In the 1940's, when 43 human menstruation was poorly understood, Professor C. J. van der Horst of the University of 44 Witwatersrand proposed using elephant shrews as a model.MANUSCRIPT To this end, he published an impressive body 45 of research. In the meantime, a breeding colony of rhesus macaques (Macaca mulatta ) had been 46 established at the Carnegie Institution of Washington [5]. It was used to give a detailed description of the 47 changes in the endometrium during the menstrual cycle and in pregnancy and to explore the physiological 48 basis for these events in primates [6,7]. This was considered a better model for human reproduction and 49 work on menstruation in elephant shrews was not pursued after van der Horst’s death. 50 In addition to describing reproduction in the Eastern rock elephant shrew (Elephantulus myurus ), van der 51 Horst made significant observations on the early development and placentation of South African mammals. 52 His work was extendedACCEPTED by scientists working on his extensive collection. These contributions are reviewed 53 below and his work on elephant shrews discussed in the light of renewed interest in the decidualization 54 process and its role in the evolution of placentation in eutherian mammals [8-10]. 55 2. Brief Biography 4 ACCEPTED MANUSCRIPT 56 Cornelius Jan van der Horst was born 11 May 1899 in Nieuwer-Amstel (now Amstelveen). He studied 57 botany and zoology at the University of Amsterdam, where early influences included the botanist Hugo De 58 Fries and the zoologist Max Weber [11-16]. His doctoral thesis from 1916 was on the motor nuclei and 59 tracts of the fish brain [17]. In the same vein, he collaborated with the Swedish zoologist Nils Holmgren to 60 describe the central nervous system of the Queensland lungfish (Neoceratodus forsteri ) [18]. He studied 61 corals under the tutelage of Stanley Gardiner of Cambridge University and in 1920 participated in a marine 62 biological expedition to the Caribbean island of Curaçao. This led to publications on corals and acorn worms 63 (Hemichordata: Enteropneusta). In 1925 he became Deputy Director of the Central Institute for Brain 64 Research in Amsterdam, but in 1928 moved to Johannesburg. 65 Max Weber, who had visited South Africa in 1894-5, encouraged van der Horst’s interest in its fauna. With 66 Stanley Gardiner as intermediary, he was offered and accepted a post as senior lecturer in the zoology 67 department of the University of the Witwatersrand [19]. He succeeded to the chair of zoology 4 years later. 68 At Johannesburg van der Horst continued his work in marineMANUSCRIPT biology, taking students on field trips to 69 Inhaca Island off the coast of Mozambique (Fig. 1) and publishing a weighty tome on the hemichordates 70 [20]. Yet soon he became intrigued by the unique mammalian fauna of South Africa and this led him to 71 study their reproduction. 72 George H. Findlay, in an affectionate portrait of his mentor, wrote that van der Horst “looked like a sedate, 73 benign Hollander, who might be seen anywhere sitting quietly in a church pew” [14]. He was far from 74 talkative and “students surmised he had lost the habit of speaking through being surrounded at home by a 75 large and purely female family!” [14]. Van der Horst had married Katherina Jacoba Aletta Vergouwen in 76 1918 and they had fourACCEPTED daughters. C. J. van der Horst died on 10 October 1951 at the age of 62. 77 3. Reproduction in the elephant shrew 78 C. J. van der Horst penned as many as 35 papers [21] on reproduction in the Eastern rock elephant shrew 79 (Elephantulus myurus jamesoni ). Many were written in collaboration with Joseph Gillman (1907-1981) 5 ACCEPTED MANUSCRIPT 80 mentor of the anthropologist Phillip V. Tobias (1925-2012) and Nobel laureate Sydney Brenner (1927- 81 present) [22]. Perhaps due to their exhaustive nature, there has been little follow up on these studies other 82 than a paper by Tripp [21] and separate studies of placentation. Only the salient features will be mentioned 83 here. 84 3.1. Ovulation and corpus luteum formation 85 E. myurus can carry one fetus in each horn of the uterus. Therefore it is remarkable that many more ova are 86 discharged from each ovary; van der Horst and Gillman suggested as many as 60 [23] and Tripp calculated a 87 mean of 49 [21].