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VU Research Portal Evolution of linoleic acid biosynthesis in Collembola and different species of arthropods Malcicka, M. 2018 document version Publisher's PDF, also known as Version of record Link to publication in VU Research Portal citation for published version (APA) Malcicka, M. (2018). Evolution of linoleic acid biosynthesis in Collembola and different species of arthropods. General rights Copyright and moral rights for the publications made accessible in the public portal are retained by the authors and/or other copyright owners and it is a condition of accessing publications that users recognise and abide by the legal requirements associated with these rights. • Users may download and print one copy of any publication from the public portal for the purpose of private study or research. • 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 the publication in the public portal ? Take down policy If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim. E-mail address: [email protected] Download date: 01. Oct. 2021 Evolution of linoleic acid biosynthesis in Collembola and different species of arthropods Miriama Malcicka VRIJE UNIVERSITEIT Evolution of linoleic acid biosynthesis in Collembola and different species of arthropods ACADEMISCH PROEFSCHRIFT ter verkrijging van de graad Doctor of Philosophy aan de Vrije Universiteit Amsterdam, op gezag van de rector magnificus prof.dr. V. Subramaniam, in het openbaar te verdedigen ten overstaan van de promotiecommissie van de Faculteit der Bètawetenschappen op dinsdag 15 mei 2018 om 11.45 uur in de aula van de universiteit, De Boelelaan 1105 door Miriama Malcicka geboren te Michalovce, Slowakije i promotor: prof.dr. J. Ellers copromotor: prof.dr. M.P. Berg ii Contents Evolution of linoleic acid biosynthesis in Collembola and different species of arthropods ....... i Contents ................................................................................................................................... iii 1. Introduction ........................................................................................................................ 1 2. An evolutionary perspective on linoleic acid synthesis in animals ..................................... 12 3. Ecomorphological adaptation in Collembola in relation to feeding strategies and habitat . 33 4. De novo synthesis of linoleic acid in multiple Collembola species ..................................... 56 5. Feeding preference and spermatophore choice of Collembola in relation to dietary linoleic acid content .............................................................................................................................. 71 6. Biosynthesis of linoleic acid in different species of Arthropoda ......................................... 84 7. Functional characterization of a Δ12 desaturase gene from Nasonia vitripennis ................ 99 8. Discussion .......................................................................................................................... 112 Bibliography .......................................................................................................................... 123 Summary ................................................................................................................................ 145 Samenvatting.......................................................................................................................... 147 Acknowledgment ................................................................................................................... 149 Curriculum Vitae ................................................................................................................... 150 Publications ............................................................................................................................ 151 Affiliation of co-authors ........................................................................................................ 154 iii 1. Introduction The role of adaptation and co-opted traits in animal evolution An important goal of evolution studies is to understand the process by which the current biodiversity has arisen and how species evolve during this process. It is often argued that species diversity is associated with the continuous growth of trait complexity in species evolution. This theory foundation was laid by J.B. Lamarck (1801), C.R. Darwin (1859) and other scientists in the 19th century. In 1835, Darwin’s crew of the ship the Beagle arrived at the Galapagos archipelago where they collected samples of plants and animals. After returning to London, Darwin noted that he had thirteen kinds of very similar sparrow, species of Geospizinae, from different islands which varied in size and shape of the beak. In his diary, popularly known as the book with the title The Voyage of the Beagle, he wrote in an amazed way: "It seems to people that from the originally small numbers of birds on this archipelago somebody chose a single species and adapted it for different purposes". Darwin described here almost exactly the principle of an evolutionary process called adaptive radiation, which is considered the precursor to the emergence of new species with the key components, adaptations to local conditions. Many complex structures in living organisms evolve as an immediately useful adaptation to one purpose, while some features could additionally gain important functions/adaptation in the later process of species’ evolution; it means they can be secondarily adaptive. For instance, if we observe the presence of a highly complex structure in an organism, which random occurrence is highly unlikely, we usually tend to explain their origin by natural selection and we seek their biological function. Even when we find that the structure is advantageous for the organism, it does not automatically mean that it has been evolved by the assumed selective pressure. In many cases, it may be co-opted, and in other cases it is not related to the natural selection at all (Grantham, 2004). A co-opted trait is a characteristic of an organism that enhances its fitness in its present role but did not evolve for that specific role by natural selection (Gould and Vrba, 1982; McLennan, 2008). Actually, the emergence of many adaptive traits by natural selection was not a direct process. Many biological structures or patterns of behavior have arisen as a result of other selection pressures than those inferred from their current biological function, and in some cases the origin of the 2 adaptive structures was not done by selective pressure (Gould and Vrba, 1982; McLennan, 2008). For instance, insect wings have probably originated from structures that originally served the process of breathing or thermoregulation (May, 1979). Therefore, insect wings originated and were formed for a long time by the selection pressures resulting from their original function. Similarly, bird wings were formed and for a long time evolved as a thermoregulation organ of some reptile groups, and only after reaching a certain developmental stage it could begin to perform its aerodynamic function and its further evolution could begin to be influenced by the selection pressures resulting from the wing function during flight (May, 1979). Another example of a co-opted trait is the exoskeletons of aquatic insects. In the aquatic environment, the exoskeleton was mainly used for the attachment of muscles, while terrestrial insect use the exoskeleton for survival on land as it is necessary to protect itself from desiccation (Zrzavý and Řičánková, 2004). So besides its primary function, the exoskeleton can also be considered as a preadaptation for life outside the aquatic environment (Zrzavý and Řičánková, 2004). The importance of pre-adaptation or co-opted trait lies in the fact that it allows evolution to proceed more quickly and in specific direction. It is not necessary to select countless new structures for each change of environment since in some cases it is possible to use structures that already exist and what is sufficient also in terms of a new function (Flegr, 1998; Zrzavý and Řičánková, 2004). Therefore, when we are studying the evolution of any biological structure, it is always necessary to think about co-option of some traits. It is important to consider not only the present biological function, but also the functions with which this structure would influence the processes of its evolutionary shaping. So, traits can be directly selected for or are co-opted. Once a lineage becomes fixed for a certain trait, descendants of that lineage will all have the derived trait unless there is a subsequent evolutionary change in that trait. This change in trait expression might result in the contrast of a trait to the ancestral trait or it might lead to completely loss of this trait (Baum, 2008). The loss of a trait can bring harmful consequences for species evolution if trait loss has fitness consequences and this essential trait is not regained or its function is not compensated for by environmental factors or a biological partner (Ellers et al., 2012; Visser et al., 2010). For instance, the loss of biosynthesis of vitamins belonging to B group in human 3 is compensated by their dietary intake from human environment (Kennedy, 2016). Thus, ecological interactions