Structure, Function and Evolution of the Labral and Frontal Glands in Termites Valeria Danae Palma Onetto
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Structure, function and evolution of the labral and frontal glands in termites Valeria Danae Palma Onetto To cite this version: Valeria Danae Palma Onetto. Structure, function and evolution of the labral and frontal glands in termites. Populations and Evolution [q-bio.PE]. Université Sorbonne Paris Cité, 2019. English. NNT : 2019USPCD027. tel-03033808 HAL Id: tel-03033808 https://tel.archives-ouvertes.fr/tel-03033808 Submitted on 1 Dec 2020 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. UNIVERSITÉ PARIS 13, SORBONNE PARIS CITÉ ECOLE DOCTORALE GALILEÉ THESE présentée pour l’obtention du grade de DOCTEUR DE L’UNIVERSITE PARIS 13 Spécialité: Ethologie Structure, function and evolution Defensiveof the labral exocrine and glandsfrontal glandsin termites in termites Présentée par Valeria Palma–Onetto Sous la direction de: David Sillam–Dussès et Jan Šobotník Soutenue publiquement le 28 janvier 2019 JURY Maria Cristina Lorenzi Professeur, Université Paris 13 Présidente du jury Renate Radek Professeur, Université Libre de Berlin Rapporteur Yves Roisin Professeur, Université Libre de Bruxelles Rapporteur David Sillam–Dussès Maitre de conférences, Université Paris 13 Directeur de thèse Jan Šobotník Chargé de Recherche, Czech University of Life Sciences Directeur de thèse Laboratoire d’Ethologie Expérimentale et Comparée Structure, function and evolution of the defensive exocrine glands in termites 2 Structure, function and evolution of the defensive exocrine glands in termites Valeria Palma Onetto A collaboration between the Termites Research Team at the Czech University of Life Sciences and the Laboratoire d’Ethologie Expérimentale et Comparée at the Université Paris 13. Structure, function and evolution of the defensive exocrine glands in termites 3 Dédicace I dedicate this work to all that people who have been my support during these hard years. To my friends for hearing my sorrows and laments. To those colleagues who without knowing me much have taken the time to speak and support me: Aleš, Cecilia and Rebeca. And, those others who became closer and provided extense conversations, knowledge and confidence: Eliska, Katka and Tomáš. I would like to thanks especially to David, my supervisor, for always providing nice words, advice, constructive criticism and all the tools I may have needed during my PhD. And finally, to the person without who it would have been impossible, to my girlfriend: Anais. Who got me up in the most difficult moments, heard all my sorrows, read my e–mails when I was not strong enough to do it by myself, encouraged me to follow my objectives and gave me the biggest reason to continue on it. My parents will never read it, they do not even understand English, but I still want to say: I am sorry. I am sorry for not have been there in these years, where things were not easy for you. I am sorry for let my sadness and pressure overcome my feelings and not had taken the first available flight when it was needed. I am sorry for letting you alone when you needed where to hold. This work is also for you, although it does not replace those moments I have missed. Contents 4 Contents CONTENTS ........................................................................................................................................................4 1. GENERAL INTRODUCTION .............................................................................................................................7 1.1 EUSOCIAL ORGANISMS ......................................................................................................................................... 8 1.2 THE TERMITES .................................................................................................................................................. 11 Phylogeny ..................................................................................................................................................... 13 Economical impact ........................................................................................................................................ 14 Termites’ abundance .................................................................................................................................... 15 1.3 DEFENSE MECHANISMS OF TERMITES .................................................................................................................... 16 The nest ........................................................................................................................................................ 16 The soldiers ................................................................................................................................................... 17 Defensive strategies in other castes ............................................................................................................. 19 1.4 CHEMICAL DEFENSES ......................................................................................................................................... 20 Exocrine glands in termites ........................................................................................................................... 21 1.5 MOTIVATION AND OBJECTIVES OF MY THESIS ......................................................................................................... 24 2. GENERAL METHODS .................................................................................................................................... 26 2.1 ANIMALS OF STUDY ........................................................................................................................................... 27 2.2 HISTOLOGY ..................................................................................................................................................... 27 2.3 MICROSCOPY .................................................................................................................................................. 27 2.4 BEHAVIOURAL TEST ........................................................................................................................................... 28 2.5 OTHERS .......................................................................................................................................................... 28 PAPER 1: THE LABRAL GLAND IN TERMITE SOLDIERS ...................................................................................... 29 INTRODUCTION ...................................................................................................................................................... 32 MATERIAL AND METHODS ....................................................................................................................................... 33 Direct Observations ...................................................................................................................................... 33 Optical microscopy and transmission electron microscopy .......................................................................... 33 Histology ....................................................................................................................................................... 33 Electron Microscopy ...................................................................................................................................... 33 Evolution of the hyaline tip ........................................................................................................................... 34 RESULTS ............................................................................................................................................................... 34 Scanning electron microscopy ...................................................................................................................... 35 Optical microscopy........................................................................................................................................ 36 Transmission electron microscopy ................................................................................................................ 36 DISCUSSION .......................................................................................................................................................... 39 CONCLUSION AND FURTHER HYPOTHESES .................................................................................................................... 42 ACKNOWLEDGEMENTS ............................................................................................................................................ 42 SUPPORTING INFORMATION ..................................................................................................................................... 43 REFERENCES .......................................................................................................................................................... 48 PAPER 2: THE LABRAL GLAND IN TERMITES: EVOLUTION AND FUNCTION .....................................................