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SPOLNI DIMORFIZEM MREŽNICE OBADA Tabanus Bromius

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SPOLNI DIMORFIZEM MREŽNICE OBADA Tabanus Bromius UNIVERZA V LJUBLJANI BIOTEHNIŠKA FAKULTETA ŠTUDIJ MOLEKULSKE IN FUNKCIONALNE BIOLOGIJE Rok JANŽA SPOLNI DIMORFIZEM MREŽNICE OBADA Tabanus bromius MAGISTRSKO DELO Magistrski študij – 2. stopnja Ljubljana, 2018 UNIVERZA V LJUBLJANI BIOTEHNIŠKA FAKULTETA ŠTUDIJ MOLEKULSKE IN FUNKCIONALNE BIOLOGIJE Rok JANŽA SPOLNI DIMORFIZEM MREŽNICE OBADA Tabanus bromius MAGISTRSKO DELO (Magistrski študij – 2. stopnja) SEXUAL DIMORPHISM OF RETINA IN THE HORSEFLY Tabanus bromius M. SC. THESIS (Master Study Programmes) Ljubljana, 2018 II Janža R. Spolni dimorfizem mrežnice obada Tabanus bromius Mag. delo. Ljubljana, Univ. v Ljubljani, Biotehniška fakulteta, Študij molekulske in funkcionalne biologije, 2018 Magistrsko delo je zaključek študija na magistrskem študijskem programu Molekulska in funkcionalna biologija. Opravljeno je bilo na Katedri za fiziologijo, antropologijo in etologijo Oddelka za biologijo, Biotehniške fakultete, Univerze v Ljubljani. Komisija Oddelka za biologijo za študij 1. in 2. stopnje je dne 17. 3. 2017 odobrila naslov magistrske naloge. Za mentorja magistrskega dela je bil imenovan izr. prof. dr. Gregor Belušič, za somentorja asist. dr. Aleš Škorjanc, za recenzenta pa prof. dr. Marko Kreft. Komisija za oceno in zagovor: Predsednik: prof. dr. Janko BOŽIČ Univerza v Ljubljani, Biotehniška fakulteta, Oddelek za biologijo Član: izr. prof. dr. Gregor BELUŠIČ Univerza v Ljubljani, Biotehniška fakulteta, Oddelek za biologijo Član: asist. dr. Aleš ŠKORJANC Univerza v Ljubljani, Biotehniška fakulteta, Oddelek za biologijo Član: prof. dr. Marko KREFT Univerza v Ljubljani, Biotehniška fakulteta, Oddelek za biologijo Datum zagovora: 31. 8. 2018 Rok Janža III Janža R. Spolni dimorfizem mrežnice obada Tabanus bromius Mag. delo. Ljubljana, Univ. v Ljubljani, Biotehniška fakulteta, Študij molekulske in funkcionalne biologije, 2018 KLJUČNA DOKUMENTACIJSKA INFORMACIJA ŠD Du2 DK UDK 591.1:595.772(043.2) KG navadni obad, spolni dimorfizem, akutna cona, mrežnica, periferni receptorji, informacijski prenos AV JANŽA, Rok, dipl. biol. (UN) SA BELUŠIČ, Gregor (mentor), ŠKORJANC, Aleš (somentor), KREFT, Marko (recenzent) KZ SI-1000 Ljubljana, Jamnikarjeva 101 ZA Univerza v Ljubljani, Biotehniška fakulteta, Študijski program Molekulska in funkcionalna biologija LI 2018 IN SPOLNI DIMORFIZEM MREŽNICE OBADA Tabanus bromius TD Magistrsko delo (Magistrski študij – 2. stopnja) OP X, 76 str., 2 preg., 45 sl., 56 vir. IJ sl JI sl/en AI Navadni obad (Tabanus bromius, Linnaeus 1758) uporablja vid za pomembne vedenjske naloge. Oba spola iščeta vodo, ki je ključna za razmnoževanje vrste in rastline, s katerih pijeta sokove. Poleg tega samci uporabljajo vid za zaznavo samic na nebu, samice pa zaznavajo plen na kompleksnem zelenem ozadju. Sestavljene oči odlikuje izrazit spolni dimorfizem. Raziskali smo anatomske in funkcionalne lastnosti fotoreceptorjev v mrežnici obeh spolov. Uporabili smo svetlobno in elektronsko mikroskopijo ter elektrofiziološke meritve. Akutni coni pri obeh spolih sta visoko specializirani za detekcijo potencialnega partnerja oz. plena. Pri samcih je akutna cona (t.i. zalezovalnik, angl. love spot) nameščena dorzofrontalno, pri samicah pa frontalno. Premer faset v akutni coni je pri samcih ≤50 µm, pri samicah pa ≤40 µm. Interomatidijski kot v akutni coni obeh spolov znaša ~1,1°. Rabdom je pri samcih dolg 350 µm, pri samicah 250 µm. Pri samicah in v ventralni mrežnici samcev sta centralni celici R7 in R8 optično sklopljeni v centralno rabdomero. Pri samcih je v zalezovalniku distalna celica (R7) pretvorjena v periferno, proksimalna celica (R8) pa je reducirana. R1-7 samcev imajo izboljšano razmerje signal/šum, razširjeno dinamično delovno območje receptorskega potenciala in vrh spektralne občutljivosti v UV in modrem (max=350,450 nm). Drugje imajo R1-6 vrh občutljivosti v zelenem (max=520 nm). Stopitvene frekvence v akutnih conah nad 30 °C dosegajo 300 Hz (samci) oz. 450 Hz (samice). Informacijska kapaciteta samčjih perifernih receptorjev je dvakrat višja kot pri samičjih (400 proti 200 bitov/s). Samci imajo v akutni coni razvit akromatski vid, optimiziran na zaznavo minimalnih zatemnitev. Njihovi fotoreceptorji dosegajo visoko informacijsko kapaciteto tako na račun izboljšanega razmerja signal/šum, kot tudi pasovne širine odziva. IV Janža R. Spolni dimorfizem mrežnice obada Tabanus bromius Mag. delo. Ljubljana, Univ. v Ljubljani, Biotehniška fakulteta, Študij molekulske in funkcionalne biologije, 2018 KEY WORDS DOCUMENTATION DN Du2 DC UDC 591.1:595.772(043.2) CX horsefly, sexual dimorphism, acute zone, retina, peripheral receptors, information transfer AU JANŽA, Rok AA BELUŠIČ, Gregor (supervisor), ŠKORJANC, Aleš (co-advisor), KREFT, Marko (reviewer) PP SI-1000 Ljubljana, Jamnikarjeva 101 PB University of Ljubljana, Biotechnical faculty, Master Study Programme in Molecular and Functional Biology PY 2018 TI SEXUAL DIMORPHISM OF RETINA IN THE HORSEFLY Tabanus bromius DT M. Sc. Thesis (Master Study Programmes) NO X, 76 p., 2 tbl., 45 fig., 56 ref. LA sl AL sl/en AB The band-eyed brown horsefly (Tabanus bromius, Linnaeus 1758) uses vision for important behavioral tasks. Both sexes use it to locate bodies of water, which are crucial for the reproduction and plants with which they feed. The males use vision to locate females, the females use it to locate their prey. The eyes are sexually dimorphic. We have investigated the anatomical and functional aspects of the retina of both sexes. We used light and electron microscopy and electrophysiological measurements. The acute zones of both sexes are highly specialised for their specific tasks – location of a potential mate or prey. In males, the acute zone or the so called love spot is located in the dorsofrontal region of the eye, in females in the frontal region. In the male acute zone, the diameter of the facets is ≤50 µm, in females ≤40 µm. The interommatidial angles in the acute zones of both sexes are ~1,1°. The length of the open rhabdom is 350 µm in males and 250 µm in females. In females, the central photoreceptors are optically coupled into central rhabdomere. In males, the situation is similar in the ventral region. In the love spot, the R7 receptor is functionally transformed into a peripheral receptor. In this case, the proximal cell R8 is greatly reduced. R1-7 in the acute zone of males have a superior SNR, a wider dynamic range of the receptor potential and a peak in the spectral sensitivity in blue and UV (max=350,450 nm). In R1-6 of the ventral region, the peak in the spectral sensitivity remains in the green part of the spectrum, similar to females (max=520 nm). The flicker fusion frequencies are ≤300 Hz in male and ≤450 Hz in female acute zones, at T>30 °C. The information capacity of the male peripheral photoreceptors is twice as high, as in females (400 vs. 200 bits/s). Male vision in the love spot is achromatic, insensitive to polarised light and adapted fort the detection of minimal intensity contrasts. Their photoreceptors reach a high information capacity on the account of a higher SNR, and also due to a broader frequency coding bandwidth. V Janža R. Spolni dimorfizem mrežnice obada Tabanus bromius Mag. delo. Ljubljana, Univ. v Ljubljani, Biotehniška fakulteta, Študij molekulske in funkcionalne biologije, 2018 KAZALO VSEBINE KLJUČNA DOKUMENTACIJSKA INFORMACIJA III KEY WORDS DOCUMENTATION IV KAZALO VSEBINE V KAZALO PREGLEDNIC VII KAZALO SLIK VIII OKRAJŠAVE IN SIMBOLI X 1 UVOD 1 1.1 NAMEN DELA 1 1.2 DELOVNA HIPOTEZA 1 2 PREGLED OBJAV 2 2.1 OPIS VRSTE 2 2.2 EVOLUCIJSKI IZVOR SESTAVLJENEGA OČESA DVOKRILCEV 3 2.3 SVETLOBA 4 2.4 ANATOMSKE ZNAČILNOSTI MUŠJEGA SESTAVLJENEGA OČESA 5 2.4.1 Zgradba sestavljenega očesa 5 2.4.2 Ločljivost in občutljivost 7 2.4.3 Apozicija, superpozicija in nevralna superpozicija 8 2.4.4 Regionalizacija 9 2.5 SPEKTRALNA IN POLARIZACIJSKA OBČUTLJIVOST 10 2.5.1 Spektralna občutljivost 10 2.5.2 Polarizacijska občutljivost 10 2.5.3 Polarizacijski vid obadov 11 2.6 FOTOTRANSDUKCIJSKA KASKADA 12 2.6.1 Rodopsin 12 2.6.2 Potek fototransdukcijske kaskade 14 2.6.3 Receptorski potencial 16 2.7 GLOBOKA PSEVDOPUPILA 16 2.8 ELEKTRORETINOGRAM 18 2.9 OSNOVE INFORMACIJSKE TEORIJE IN NJENA UPORABA V NEVROBIOLOGIJI 19 2.9.1 Vsebnost informacije in njena bitna narava 19 2.9.2 Informacijski prenos 20 2.10 EKONOMIKA MUŠJE FOTOTRANSDUKCIJE 22 3 MATERIAL IN METODE 23 3.1 LOVLJENJE IN PRIPRAVA ŽIVALI 23 3.2 MAKROSKOPSKI POSNETKI 23 3.3 MIKROSKOPIJA 23 3.3.1 Svetlobna mikroskopija 23 VI Janža R. Spolni dimorfizem mrežnice obada Tabanus bromius Mag. delo. Ljubljana, Univ. v Ljubljani, Biotehniška fakulteta, Študij molekulske in funkcionalne biologije, 2018 3.3.1.1 Analiza interomatidijskih kotov 23 3.3.2 Elektronska mikroskopija 24 3.3.2.1 Presevna elektronska mikroskopija 24 3.3.2.2 Vrstična elektronska mikroskopija: Analiza velikost faset 24 3.4 ELEKTROFIZIOLOGIJA 25 3.4.1 ERG 25 3.4.2 Znotrajcelični posnetki 25 3.4.2.1 Spektralna občutljivost 26 3.4.2.2 Receptorski potencial 26 3.4.2.3 Kotna frekvenca 26 3.4.2.4 Sprejemni kot 26 3.4.2.5 Meritve informacijskega prenosa 26 4 REZULTATI 29 4.1 ANATOMIJA IN ULTRASTRUKTURA 29 4.2 ELEKTROFIZIOLOŠKE MERITVE 37 5 RAZPRAVA 52 6 SKLEPI 58 7 POVZETEK 59 8 VIRI 60 ZAHVALA VII Janža R. Spolni dimorfizem mrežnice obada Tabanus bromius Mag. delo. Ljubljana, Univ. v Ljubljani, Biotehniška fakulteta, Študij molekulske in funkcionalne biologije, 2018 KAZALO PREGLEDNIC Preglednica 1: Površine posameznih rabdomer v zalezovalniku. ................................... 35 Preglednica 2: Multiple primerjave parov stopitvenih frekvenc v Anova analizi.
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