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Pycnial Nectar of Rust Fungi Induces Cap Formation on Pycniospores of Opposite Mating Type

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Pycnial Nectar of Rust Fungi Induces Cap Formation on Pycniospores of Opposite Mating Type Pycnial Nectar of Rust Fungi Induces Cap Formation on Pycniospores of Opposite Mating Type Y. Anikster; T. Eilam; L. Mittelman; L. J. Szabo; W. R. Bushnell Mycologia, Vol. 91, No. 5. (Sep. - Oct., 1999), pp. 858-870. Stable URL: http://links.jstor.org/sici?sici=0027-5514%28199909%2F10%2991%3A5%3C858%3APNORFI%3E2.0.CO%3B2-3 Mycologia is currently published by Mycological Society of America. Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/about/terms.html. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/journals/mysa.html. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. The JSTOR Archive is a trusted digital repository providing for long-term preservation and access to leading academic journals and scholarly literature from around the world. The Archive is supported by libraries, scholarly societies, publishers, and foundations. It is an initiative of JSTOR, a not-for-profit organization with a mission to help the scholarly community take advantage of advances in technology. For more information regarding JSTOR, please contact [email protected]. http://www.jstor.org Thu Feb 28 11:11:03 2008 ~~f~coloqzcr,91(i), 1999, pp 858-870 Q 1999 b\ The hhcolog~calSoc~eti of I~nrr~c'~,1'iwrrnce. IS 66044-8897 Pycnial nectar of rust fungi induces cap formation on pycniospores of opposite mating type1 Y Anikster presence of polysaccharides or glycoproteins contain- T. Eilam ing N-acetylglucosamine. The cap-inducing activity of Department of Botany and Institute for nectar was lost if the nectar was boiled or autoclaved. Cereal Crops Improvement, Tel Auiz~CTniuersity, Cap-inducing nectar contained a complex of high Ramat Auiu, 69978, Israel molecular weight proteins larger than 100 kDa as L. Mittelman shown by native polyacrylamide gel electrophoresis Sackler School of Medicine, Tel Auiv University, analysis. This protein conlplex had cap-inducing ac- Ramat Auiu, 69978, Israel tivity as determined by placing pycniospores directly on gels after electrophoresis. In SDS gels, 6-7 poly- L. J. Szabo peptides ranging in size from 14-70 kDa were ob- W. R. Bushnel12 served, but bioassays of these polypeptides for cap USDA-ART, Cereal Disease I,aboratory, 1551 Lindig Street, C'niuersity of Minnesota, St. Paul, induction were negative. The results indicate that Minnesota 55108, USA pycnial nectar of several rust species contains high molecular weight cap-inducing proteins which are mating type-specific and induce pycniospore cap for- Abstract: Pycnial nectar (including pycniospores) mation as an early event associated with processes transferred between pycnia of opposite mating type leading to fertilization. (as indicated by subsequent aecium formation) in- Key Words: aecia, aecium, fertilization, mating duced formation of a cap on one end of pycnio- type, pycnia, pycnium, Uredinales spores. The polar caps developed with seven species of Puccinia and four of Uromyces, but not with P he- lianthi, Tranzschelia pruni-spinosae, or ll vignae. Caps were induced equally in reciprocal transfers of nectar between pycnia of opposite mating type. The Mating in rust fungi (Uredinales) takes place in hap- caps stained with India ink, or labeled with colloidal loid pycnia produced on pycnial host plants. In spe- gold or wheat germ agglutinin conjugated with fluo- cies investigated, pycniospores produced in pycnia of rescein isothiocyanate (WGA-FITC), but were not vis- one mating type fertilize pycnia of a second mating ible in unstained preparations. Cap formation started type by fusion to a receptive hypha, transfer of a nu- within 10 min of nectar transfer and was completed cleus into the hypha, followed in most rust fungi by in 20-60 min. Nectar retained cap-inducing activity formation of dikaryotic hyphae that produce aecia after pycniospores were removed by centrifugation, (Allen 1933, Craigie 1927, 1931, 1933, Pierson 1933). whereas pycniospores washed free of nectar did not The pycniospores are produced in pycnial nectar, a induce caps. Pycniospores that were removed from a solution of high sugar concentration thought to en- pycnium and induced to form caps by pycniospore- courage visits by insects, which then carry nectar and free nectar of opposite mating type did not induce pycniospores from one pycnial cluster to another. In aecia when returned to the original pycnium, show- a comprehensive investigation of fertilization in rust ing that cap formation alone was not sufficient for fungi, Buller (1950) described formation of fusion completion of mating processes. Caps were removed tubes from the receptive hyphae. The tube grows to- by treatment with proteinase K, sodium dodecyl sul- ward a pycniospore, which in turn, produces a very fate (SDS), or 0.01 N HCl, indicating the presence short tiny papilla where fusion then takes place. Bull- of protein. Labeling by WGA-FITC suggested the er (1950) suggested that stimuli act at a distance to induce directed fusion tube growth and papilla for- Accepted for publicatiori April 13, 1999. mation. Since Buller (1950), little has been added to ' Paper So. 981220036 of the Scientific Journal Series of thr hlin- knowledge of the mode of attraction or details of the nesota i\gric~~lturalExpel-iment Station. Mention of n traticrliark fusion process. Likewise, the possible role of nectar rlarne or proprietarx product does not constitute a guarantee of Tel kiv Llli\.ersit!. the Ynivrrsin of hlinnesota, or the V.S. De- in fertilization processes other than as an attractant par-tment of Agr-ic111r11rc. for insect vectors has not been investigated. ' Corresponding author. email: billb@pi~ccini.crl.umn.rrlu As part of a project to learn more about fertiliza- T.-\RI.EI. Species of Puccinia, Tranzschelia and Uromyces tested for nectar-induced pycniospore caps S~ecies Collectiona Telial host Pvcnial host Pziccinia allii (DC.) Rud 8782, 8827, 8837 Allizcm ampeloprasum L. Allium ampelqbrasum l? c~ronataCorda 8789, 8805, 8822 Avena sativa I>. Rhamnus palaestinus Boiss. I? grnminis Pers. f. sp. tritici 9522 Triticum aestitium L. Berberis vulgnris L. Eriks. & E. Henn. I? pminis f. sp. secalis Eriks. 2969 Agropyron r~pens(L.) Pal. 3. vulgaris 8s E. Henn. l? helianthi Schwein. 2971h, 2972", 2973"; Helitznthus annltus L. Heliantlzus annuus L. 2974h, 8784, 8785, 8829 I? hordui G. Otth 1817, 1827, 1836, 1837 Hordeum spontaneum C. Ornithogalum spp. &ch. I? recondita Roberge ex Des- 9310, 9567, 9569 Aeg~lqbslongzssinza Schweinf. Anchusa aggregata Lehm. maz er Muschler emend Eig. l? recondita 9579, 9.586, 9910 AP. ovata I,. Echium glom~ratumPoiret l? recondita 2889< Srcalv cereale L. Lycopsis arvensis L. l? reconditn 77024, 77042 Triticum turgzdum ssp. durum An. italica Retz. (Desf.) MK. I? sorghi Schwein. 8790, 8794 Zen mays L. Oxalis corniculata L. l? triticina Erikss." 77025, 9675, 9583 7: aestivum Thalictrurn speciosissimz~mI,. Tranzschelia pruni-spinosae 8904 Prz~nusamygdalus Batsch. Anemone colanarin L. (Pers.) Dietel LTromyces hippomarathricola S. 8905 Bilacunaria boissieri Pimenov Bilacunarin boissieri da Cam et Tichomirov I: hordeastri Guyot 6422, 6437 Hordeurn lrulbosum L. Scilla autumnalis L. CT. horduinus (Arth.) Barth 77064h, 77065" H. pusillum Nutt. Nothoscordium bivalve (L.) Britt. L! reichrrtii Anikst. & MTahl 6385 H. hulbosum Scilla Izyacinthoides L. U vi<gna~Barclay CPR-1' Vicpa unguiculata L. Vigna unpiculata Collections were from Israel, except as noted. Collections are as defined by Anikster et a1 (1997) to include aecial, pycnial, telial and uredinial progeny from selfing original teliospore collections. From USA. ' From Czech Republic (supplied by Z. Urban). I' l? tn'ticina designates leaf rust of wheat (following Savile 1981) which is distinct from l? recondita, which includes leaf rusts of rye and wild wheats (Anikster et a1 1997). From USA (supplied by I(. Mendgen). tion in rust fungi, we tested a number of staining recondita, collections were of four types differing in range procedures to help visualize fusion processes. Certain of both telial and pycnial hosts (TABLFI) (hikster et a1 stains revealed that a cap was produced on one end 1997). The most abundant nectar was obtained with l? re- of pvcniospores when they were treated with nectar condita collections from Anchusn italica and Echium glom- eraturn. of opposite mating type. Here, we describe cap for- Pycnia were obtained in the greenhouse by inoculating mation, demonstrate the correlation between cap pycnial host species with basidiospores from germinating formation and nectar mating type, and give prelimi- teliospores following procedures of Anikster (1986) and An- nary evidence that the cap-inducing substances in ikster et al (1997). If pycnial clusters originating from a nectar are proteins. single basidiospore were needed, plants were inoculated with low numbers of basidiospores. If nectar from multiple pycnial clusters was to be combined, plants were inoculated with higher numbers of basidiospores. To maximize volume Nectar and pycniospores were obtained from pycnial hosts of nectar, plants were placed in a cylindrical, covered, hu- infected by 14 rust species (TABLEI). For 10 of the rust midity chamber at the time pycnia brgan to develop, 8-15 species, pycnia were obtained with more than one collec- d after inoculation. For experiments on cap induction with- tion (T.\RI.E1) to increase the likelihood that a collection in pycnial clusters, plants with pycnia were transferred from giving abundant nectar would be found. In some cases, nec- the greenhouse to a plant growth chamber (Karl Weiss) at tar from two or three collections was combined to obtain 16 t 2 C and a daily 12-h photoperiod.
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