2
2014. X, 299 pages, 233 mostly coloured figures, 4 tables, 17 x 24 cm, hardcover ISBN 978-3-510-65287-7 39.80 € www.schweizerbart.de/9783510652877 This book presents a contemporary and imagina- tive synopsis of numerous biological aspects of cilia/Àagella and ciliates/Àagellates. It comprises contributions by a dozen renowned experts from all over the world, which summarize our current un- derstanding, essentially the results obtained and progress made during the last ¿ve decades of re- search of cilia/Àagella ultrastructure, cell biology, organellar function, motility, taxonomy/systematics, symbiosis, and biodiversity. It provides numerous suggestions for future re- search. The volume is lavishly illustrated by numerous line drawings and light- and electron microscopic images. It addresses advanced students of biology and zoology, and all scientists teaching and working in cell biology and protistology. Schweizerbart Science Publishers Johannesstr. 3 A, 70176 Stuttgart, Germany. Tel. +49 (711) 351456-0 Fax. +49 (711) 351456-99 [email protected] www.borntraeger-cramer.de E Cilia and Flagella Ciliates and Flagellates Ultrastructure and cell biology, function and systematics, symbiosis and biodiversity Klaus Hausmann Renate Radek (Eds.) E Binary fission of the ciliate Didinium nasutum. Magn. 700x (photo: Juliane Tröger, Berlin, Germany). My first attempt was to investigate the di- gestion of Didinium. Lysosomes had recently been discovered by de Duve (1963), and the little bit I knew about Didiniumsuggested to me that they were very inefficient feeders/di- gesters. So I imagined that Didinium might utilize the labialization of Paramecium’s own lysosomes to digest Paramecium; a project I believed worthy of investigation. If you want to get some idea of what has transpired over the years, compare my original protozoology textbook (Mckinnon and Hawes, 1961) with today’s (Hausmann et al., 2003). The beginning The natural history of Didiniumand its rela- tion with Paramecium was well described by Balbiani (1873), Thon (1905), Jennings (1906), Mast (1909), Calkins (1915), Beers (1925), and others. Each remarked on the powerful swimming, the remarkably consis- tent pattern of spiral turning as didinia swam and engaged an object, backed slightly, always turning to the right and rapidly swimming straight ahead again spiraling as it went. If di- dinia encountered suitable prey, the organisms engaged with their to Thon) or and engulfed the prey in a matter of a minute or two, regardless of the size of the prey. Not infrequently, the prey was larger than the predator. In fact, some claimed the prey could be many times larger than the Didinium. Hausmann/Radek (eds): Cilia/Flagella – Ciliates/Flagellates © Schweizerbart Science Publishers Stuttgart, 2014 www.schweizerbart.de/9783510652877 Cellular Architecture, Growth, Morphogenesis, Chemoattractants, and Loose Ends by Gregory A. Antipa That first impression of Parameciumin my high school biology class sowed the seed for my career-spanning focus on microscopy and protists. Little did I suspect that this simple introduction would inspire my first wonderment of How do they do that? After complet- ing a bachelor’s degree in cell biology, my career path came together in September of 1963. As crates containing an electron microscope, a JEM T6S, were de- livered to Science Hall, I entered graduate school. The microscope and I rarely parted. Soon we were delving into the mysteries of Didinium. Transmission electron microscopes (TEM) were bringing much new information into cell biology and protistology. Fig.1: Caught in the act: Didiniumcaptures and ingests / Hausmann/Radek Hausmann/Radek Hausmann/Radekl / l ll l ll ll ll t h b © Schweizerbar © Schweizerbar h 4 www.schweizerbCilia and Flagella – Ciliates and Flagellates 5 Fig. 4: Cilium in transverse section (a) and in combined schematic 9 x 2 + 2 representation (b). Three-dimen- sional reconstruction of the axoneme (ce) and a microtubular doublet (f). A-tub A-microtubule, B-tub B-micro- tubule, cmt central pair of microtubules, dy dynein arm, pm plasma membrane, rsp radial spoke (after Haus- mann and Gradias). Magn. 200,000x. Fig. 5: Ciliary base. a Longitudinal section of axoneme (axn) and kinetosome (ks). b Transverse section at the transitional region between axoneme and kinetosome; nine axonemal doublets (upper part) and beginning of nine kinetosomal triplets (lower part). c Nine fully developed kinetosomal triplets per kinetosome. dSchematic representation (after Dentler). Magn. a–c 60,000x. seem to beat in a two-dimensional plane (Fig. 3). However, more detailed investigations re- veal that cilia also beat in a helical or three- dimensional manner, and that some flagella move only in a single plane. Axonemal structure Cross sections of cilia and flagella show the axoneme with its characteristic arrangement of microtubules (the so-called 9 × 2 + 2 pat- tern), which is surrounded by the plasma membrane (Fig. 4). There are nine peripheral double microtubules (i. e. doublets) and two individual central microtubules (i. e. singlets). The doublets each consist of a complete mi- crotubule with 13 protofilaments (A-tubule) and an attached microtubule (B-tubule) which has only ten protofilaments of its own but shares three protofilaments with the A-tubule. Each A-tubule is the origin of pairs of arm- like structures (dynein arms) which are di- rected toward the B-tubule of the adjacent doublet. The doublets are interconnected by nexin links which also attach the A-tubule of each doublet to the B-tubule of the adjacent doublet. The central microtubules are sur- rounded by central sheath material from which radial spokes arise; these spokes are directed towards and connected with the A-tubules of each doublet. None of these structures is con- tinuous along the length of the axoneme, and each has its own periodicity of occurrence. Structure of basal bodies (kinetosomes) and axonemal tips The organization of the basal body differs from that of the axoneme. The central tubules of the cilium terminate at an axial granule; the peripheral doublets continue into the basal body where an additional C-tubule is added. Like the B-tubule, the C-tubule is an incom- plete microtubule, which in this case shares protofilaments with the B-tubule (Fig. 5). Cili- ates often have non-ciliated (sometimes re- ferred to as barren) basal bodies which may be paired with ciliated ones. The basal bodies of flagellates and ciliates are associated with mi- crotubular and filamentous structures which are arranged in a species-specific manner. The number, site of insertion, and organiza- tion of these elements may be used to deter- mine phylogenetic relationships between dif- ferent groups of ciliates and flagellates (see the chapter by Denis H. Lynn, this book). Since all cilia/flagella have a common ori- gin, it is likely that some of the associated root structures are homologous as well. This may help to identify homologous flagella in differ- 4 K. Hausmann and R. Radek Fig. 2: Cilia of ciliates in front view (a, Homalozoon) and in longitudinal section (b, Paramecium). axn axo- neme, bb basal body (= kinetosome), pm plasma membrane. Magn. a 1,700x, b 45,000x. Fig. 3: Diagram illustrating undulatory flagellar (a) and ciliary movements (b) (after Satir). sample pages

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Page 1: Cilia and Flagella Ciliates and Flagellates

2014. X, 299 pages, 233 mostly coloured figures, 4 tables,17 x 24 cm, hardcover

ISBN 978-3-510-65287-7 39.80 € www.schweizerbart.de/9783510652877

This book presents a contemporary and imagina-tive synopsis of numerous biological aspects of cilia/ agella and ciliates/ agellates. It comprises contributions by a dozen renowned experts from all over the world, which summarize our current un-derstanding, essentially the results obtained and progress made during the last ve decades of re-search of cilia/ agella ultrastructure, cell biology, organellar function, motility, taxonomy/systematics, symbiosis, and biodiversity.

It provides numerous suggestions for future re-search.

The volume is lavishly illustrated by numerous line drawings and light- and electron microscopic images.

It addresses advanced students of biology and zoology, and all scientists teaching and working in cell biology and protistology.

Schweizerbart Science PublishersJohannesstr. 3 A, 70176 Stuttgart, Germany. Tel. +49 (711) 351456-0 Fax. +49 (711) [email protected] www.borntraeger-cramer.deE

Cilia and FlagellaCiliates and FlagellatesUltrastructure and cell biology, function and systematics, symbiosis and biodiversity

Klaus Hausmann • Renate Radek (Eds.)

E

Binary fission of the ciliate Didinium nasutum. Magn. 700x(photo: Juliane Tröger, Berlin, Germany).

My first attempt was to investigate the di-gestion of Didinium. Lysosomes had recentlybeen discovered by de Duve (1963), and thelittle bit I knew about Didinium suggested tome that they were very inefficient feeders/di-gesters. So I imagined that Didinium mightutilize the labialization of Paramecium’s ownlysosomes to digest Paramecium; a project Ibelieved worthy of investigation. If you wantto get some idea of what has transpired overthe years, compare my original protozoologytextbook (Mckinnon and Hawes, 1961) withtoday’s (Hausmann et al., 2003).

The beginningThe natural history of Didinium and its rela-tion with Paramecium was well describedby Balbiani (1873), Thon (1905), Jennings(1906), Mast (1909), Calkins (1915), Beers(1925), and others. Each remarked on thepowerful swimming, the remarkably consis-tent pattern of spiral turning as didinia swamand engaged an object, backed slightly, alwaysturning to the right and rapidly swimmingstraight ahead again spiraling as it went. If di-dinia encountered suitable prey, the organismsengaged with their mittlere Strang (accordingto Thon) or seizing organ (according to Calkins)

and engulfed the prey in a matter of a minuteor two, regardless of the size of the prey. Notinfrequently, the prey was larger than thepredator. In fact, some claimed the prey couldbe many times larger than the Didinium.

Hausmann/Radek (eds): Cilia/Flagella – Ciliates/Flagellates © Schweizerbart Science Publishers Stuttgart, 2014www.schweizerbart.de/9783510652877

Cellular Architecture, Growth,Morphogenesis, Chemoattractants, and Loose Ends

byGregory A. Antipa

That first impression of Paramecium in my high school biology class sowed the seed formy career-spanning focus on microscopy and protists. Little did I suspect that this simpleintroduction would inspire my first wonderment of How do they do that? After complet-ing a bachelor’s degree in cell biology, my career path came together inSeptember of 1963. As crates containing an electron microscope, a JEM T6S, were de-livered to Science Hall, I entered graduate school. The microscope and I rarely parted.Soon we were delving into the mysteries of Didinium. Transmission electron microscopes(TEM) were bringing much new information into cell biology and protistology.

Fig.1: Caught in the act: Didinium captures and ingestsParamecium (SEM) (see also Wessenberg and Antipa,1970).to Thon) or seizing organ (according to Calkins)

/ d k d l / l ll l / l ll/ d kH /R d kHausmann/RadekHausmann/RadekHausmann/Radek ( d ) l /( d ) Cili /(eds): Cilia/(eds): Cilia/(eds): Cilia/ l ll lFl ll CilFlagella – CilFlagella – CilFlagella Cil / l lli t /Fl lliates/Flagellaiates/Flagellaiates/Flagellattestestes h b© Schweizerbar© Schweizerbar blt Science Publt Science Publ hishers Stuttgaishers Stuttga 4rt 2014rt, 2014

wwww.schweizerbart.de/9783510652877

1970).

Cilia and Flagella – Ciliates and Flagellates 5

Fig. 4: Cilium in transverse section (a) and in combined schematic 9 x 2 + 2 representation (b). Three-dimen-sional reconstruction of the axoneme (c–e) and a microtubular doublet (f). A-tub A-microtubule, B-tub B-micro-tubule, cmt central pair of microtubules, dy dynein arm, pm plasma membrane, rsp radial spoke (after Haus-mann and Gradias). Magn. 200,000x.

Fig. 5: Ciliary base. a Longitudinal section of axoneme (axn) and kinetosome (ks). b Transverse section at thetransitional region between axoneme and kinetosome; nine axonemal doublets (upper part) and beginningof nine kinetosomal triplets (lower part). c Nine fully developed kinetosomal triplets per kinetosome. d Schematicrepresentation (after Dentler). Magn. a–c 60,000x.

seem to beat in a two-dimensional plane (Fig.3). However, more detailed investigations re-veal that cilia also beat in a helical or three-dimensional manner, and that some flagellamove only in a single plane.

Axonemal structureCross sections of cilia and flagella show theaxoneme with its characteristic arrangementof microtubules (the so-called 9 × 2 + 2 pat-

tern), which is surrounded by the plasmamembrane (Fig. 4). There are nine peripheraldouble microtubules (i. e. doublets) and twoindividual central microtubules (i. e. singlets).The doublets each consist of a complete mi-crotubule with 13 protofilaments (A-tubule)and an attached microtubule (B-tubule) whichhas only ten protofilaments of its own butshares three protofilaments with the A-tubule.

Each A-tubule is the origin of pairs of arm-like structures (dynein arms) which are di-rected toward the B-tubule of the adjacentdoublet. The doublets are interconnected bynexin links which also attach the A-tubule ofeach doublet to the B-tubule of the adjacentdoublet. The central microtubules are sur-rounded by central sheath material from whichradial spokes arise; these spokes are directedtowards and connected with the A-tubules ofeach doublet. None of these structures is con-tinuous along the length of the axoneme, andeach has its own periodicity of occurrence.

Structure of basal bodies (kinetosomes)and axonemal tipsThe organization of the basal body differsfrom that of the axoneme. The central tubulesof the cilium terminate at an axial granule; theperipheral doublets continue into the basalbody where an additional C-tubule is added.Like the B-tubule, the C-tubule is an incom-plete microtubule, which in this case sharesprotofilaments with the B-tubule (Fig. 5). Cili-ates often have non-ciliated (sometimes re-ferred to as barren) basal bodies which may bepaired with ciliated ones. The basal bodies offlagellates and ciliates are associated with mi-crotubular and filamentous structures whichare arranged in a species-specific manner.The number, site of insertion, and organiza-tion of these elements may be used to deter-mine phylogenetic relationships between dif-ferent groups of ciliates and flagellates (see thechapter by Denis H. Lynn, this book).

Since all cilia/flagella have a common ori-gin, it is likely that some of the associated rootstructures are homologous as well. This mayhelp to identify homologous flagella in differ-

4 K. Hausmann and R. Radek

Fig. 2: Cilia of ciliates in front view (a, Homalozoon)and in longitudinal section (b, Paramecium). axn axo-neme, bb basal body (= kinetosome), pm plasmamembrane. Magn. a 1,700x, b 45,000x.

Fig. 3: Diagram illustrating undulatory flagellar (a)and ciliary movements (b) (after Satir).

sample pages

Page 2: Cilia and Flagella Ciliates and Flagellates

Table of contentsList of Contributors . . . . . . . . . . . . . . . . . . . VPreface . . . . . . . . . . . . . . . . . . . . . . . . . VIIIntroductionCilia and Flagella – Ciliates and FlagellatesKlaus Hausmann and Renate Radek . . . . . . . . . . 3UltrastructureCellular Architecture, Growth, Morphogenesis, Chemo-attractants, and Loose EndsGregory A. Antipa . . . . . . . . . . . . . . . . . . . . 23Ejection, Ingestion, Digestion, and Expulsion in CiliatesKlaus Hausmann . . . . . . . . . . . . . . . . . . . . 47Cell BiologyA Song of Praise for Paramecium as a Model in Vesicle Traf ckingA Sotto Voce Praise in Retrospect with Certain ReservationHelmut Plattner . . . . . . . . . . . . . . . . . . . . . 69Ciliate Mating Types and PheromonesPierangelo Luporini, Claudio Alimenti, and Adriana Vallesi . . . . . . . . . . . . . . . . . . . . . . . . . 95MotilityEncounters with CiliaMichael A. Sleigh . . . . . . . . . . . . . . . . . . . 121How do Protists keep up?Hans Machemer . . . . . . . . . . . . . . . . . . . . 133Ctenophores and Termites – Systems for MotilitySidney L. Tamm . . . . . . . . . . . . . . . . . . . . 147Taxonomy and SystematicsKinetids, Concepts, and CoincidencesDenis H. Lynn . . . . . . . . . . . . . . . . . . . . . 175

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On Algal and other Protist Flagella and CiliaØjvind Moestrup . . . . . . . . . . . . . . . . . . . . 189SymbiosisInsights into the Paramecium-Holospora and Parameci-um-Chlorella SymbiosesMasahiro Fujishima and Yuuki Kodama . . . . . . . . 203Prokaryotic Endosymbionts in CiliatesHans-Dieter Görtz . . . . . . . . . . . . . . . . . . . 229Symbionts of Symbionts – Termite Flagellates and their Bacterial AssociationsRenate Radek and Jürgen F. H. Strassert . . . . . . . 239BiodiversitySmallest Protists in the Deepest Depths – Flagellates from Abyssal Sea FloorsKlaus Hausmann . . . . . . . . . . . . . . . . . . . 255Retro- and ProspectiveFive Decades of Research in Protistology – What have we learned?Jens Boenigk . . . . . . . . . . . . . . . . . . . . . . 267AddendumCurricula Vitae of Contributors . . . . . . . . . . . . . 277Humor . . . . . . . . . . . . . . . . . . . . . . . . 287Acknowledgements . . . . . . . . . . . . . . . . . . 290Index . . . . . . . . . . . . . . . . . . . . . . . . 291

Klaus Hausmann • Renate Radek (Eds.) Cilia and Flagella / Ciliates and Flagellates

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Klaus Hausmann, Renate Radek

Festschrift 25 Jahre Deutsche Gesellschaft für Protozoologie1981–20062006. 277 Seiten, zahlr. Photos u. Abb., gebunden, 17 x 24 cmISBN 978-3-510-65217-4 19.00 €www.schweizerbart.de/9783510652174

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