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Development in Neurospora and Related Ascomycetes (Electrophoretic Variants/Serotypic Variants/Sordariaceae) JUNE BOWMAN NASRALLAH and ADRIAN M

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Development in Neurospora and Related Ascomycetes (Electrophoretic Variants/Serotypic Variants/Sordariaceae) JUNE BOWMAN NASRALLAH and ADRIAN M Proc. Natl. Acad. Sci. USA Vol. 74, No. 9, pp. 3831-3834, September 1977 Botany Occurrence of a major protein associated with fruiting body development in Neurospora and related Ascomycetes (electrophoretic variants/serotypic variants/Sordariaceae) JUNE BOWMAN NASRALLAH AND ADRIAN M. SRB Section of Genetics, Development, and Physiology, Cornell University, Ithaca, New York 14853 Contributed by Adrian M. Srb, June 20, 1977 ABSTRACT Electrophoretic and immunological analysis as strain T-220 by J. H. Warcup, and designated N. tetrasperma of fruiting body (perithecial) extracts demonstrates the occur- (Warcup); N. tetrasperma strains 1270 (85A) and 1271 (85a), rence of a major phase-specific perithecial protein in all Neu- rospora species and in the closely related Gelasinospora cerealis derived from an isolate by B. 0. Dodge, obtained from the and Sordaria fimicola. The perithecial proteins from these FGSC, and designated N. tetrasperma (Dodge); N. toroi (FGSC different species fall into a number of groups with different no. 688) obtained from the FGSC; (c) as representatives of the electrophoretic mobilities. They appear to be immunologically homothallic species-N. terricola, obtained from S. E. closely related but not identical to one another even within the Gochenaur, N. africana (Africana N200), N. dodgei (ATCC same genus, with only partial identity exhibited between the 15509), N. galapagosensis (Galapagosensis G349), and N. heterothallic and pseudohomothallic Neurospora on the one hand and the homothallic Neurospora on the other hand. In lineolata (ATCC 18966), all obtained from the FGSC. immunological analysis of fruiting body extracts of the other The Neurospora strains were selected to represent established Ascomycetes, Podospora anserina, Cochliobolus maydis, and species of the genus, essentially as classified by Frederick et al. Aspergillus nidulans, and of ascus extracts of Saccharomyces (3). Some named species were omitted, as warranted by more cerevisiae, no crossreaction with the Neurospora perithecial recent taxonomic work, namely, the exclusion of the stromatic protein was found. N. phoenix from the genus (4), or by lack of live material, as is the case for N. erythrea. A strain of N. torcn was included in the Biochemical analysis of components of the Neurospora life serological study although recent work on the crossing behavior cycle has revealed, in N. crassa, N. sitophila, N. tetrasperma, of strains and N. terricola, a specific association between fruiting body assigned to this species suggests their identity with (perithecial) development and a major protein species resolv- strains of N. tetrasperma (5, 6). able on Other Genera. Members of the other genera tested were polyacrylamide gels (1, 2). This- protein can be detected Gelasinospora cerealis, obtained from the FGSC; Podospora in unfertilized fruiting bodies, but its concentration increases anserina, from D. Marcou; Sordaria fimicola, from L. S. Olive; drastically in fertilized fruiting bodies until 4 or 5 days after Aspergillus nidulans (FGSC no. 4), from Etta Kifer; Saccha- fertilization, when it constitutes a major fraction of the total romyces cerevsiae, from G. R. Fink; and Cochliobolus maydis, perithecial proteins. Later in maturation, as perithecial contents from are lost in the perithecial exudate and subsequently in spore 0. C. Yoder. discharge, the levels of the major perithecial protein decrease Culture Techniques. Maintenance of stocks and of cultures (1, 2). Detailed reports of this type of perithecial protein have for immunological work with vegetative mycelia of Neuro- so far been limited to the usual laboratory species, N. crassa and spora, Gelasinospora, and Sordaria was at 250 on the minimal N. tetrasperma. An important biological function in fruiting medium of Beadle and Tatum (7) supplemented with 2% agar body maturation for such a phase-specific protein would be when solid medium was required. Vegetative mycelia of the indicated if a comparable protein species were found to occur homothallic strains were produced as submerged cultures in more generally among Ascomycetes. All the available Neu- Erlenmeyer flasks containing liquid minimal medium and rospora species and members of a number of other Ascomy- harvested before fruiting bodies formed. Production of proto- cetous genera have now been examined for the presence of such perithecia and crosses involving Neurospora and Gelasinospora a protein, as summarized in this report. were carried out at 250 on Difco cornmeal agar or on the liquid crossing medium of Westergaard and Mitchell (8) adjusted to MATERIALS AND METHODS pH 5.7, as described earlier (1), and supplemented with 2% agar where required. Strains of Neurospora. The strains of Neurospora used in Podospora perithecia were produced on Difco this study were the following: (a) as representatives of the cornmeal agar in the light; Sordaria crosses were made on en- heterothallic species-the standard laboratory wild-type St. riched cornmeal agar (9); Aspergillus crosses were made ac- Lawrence strains of N. crassa, 74A and 77a; Honduras 3A and cording to Pontecorvo et al. (10), and sporulating cultures of la, wild strains of N. crassa isolated in Honduras and obtained Saccharomyces were produced according to Fowell (11). Co- from R. H. Stover and S. R. Freiburg; N. intermedia P420 and chilobolus perithecia were provided by 0. C. Yoder. P405, obtained from the Fungal Genetics Stock Center (FGSC); Harvesting of Material and Biochemical and Immuno- and N. sitophila 540-34A and 2a from J. R. S. Fincham; (b) as chemical Analysis. Harvesting and extraction of samples on representatives of the pseudohomothallic species-homokar- 0.1M phosphate buffer (pH 7.0) and electrophoresis on 7.5% yotic strains of N. polyacrylamide gels were as described earlier (1). tetrasperma from Borneo, originally isolated The protein antigen for immunological studies was obtained The costs of publication of this article were defrayed in part by the from perithecial extracts of N. crassa or N. tetrasperma payment of page charges. This article must therefore be hereby marked (Dodge) which were subjected to electrophoresis. The protein "advertisement" in accordance with 18 U. S. C. §1734 solely to indicate was eluted in 0.1 M phosphate buffer (pH 7.0) from acrylamide this fact. gel slices corresponding to the protein band of interest. The gel 3831 Downloaded by guest on September 24, 2021 3832 Botany: Nasrallah and Srb Proc. Natl. Acad. Sci. USA 74 (1977) ORIGIN _N'--IA Ii I iii11 FRONT-- Hon gal Iin tern afri dod Gel ANODE FIG. 2. Electrophoretic separation of the soluble proteins of perithecial extracts. Electrophoretic variants of the major perithecial protein are shown. Abbreviations are given in the legend of Fig. 1. of Celasinospora cerealis, and of Sordaria fimicola, but not for perithecial extracts of Podospora anserina and Cochilobolus FIG. 1. Double diffusion patterns obtained with fruiting body maydis or for cleistothecial extracts of Aspergillus nidulans or extracts. Central wells contained antiserum produced against N. for ascus extracts of Saccharomyces cerevisiae. Furthermore, tetrasperma (Dodge) perithecial antigen; peripheral wells contained on the basis of the immunological reactions obtained with two fruiting body extracts. afri, N. africana; Asp, Aspergillus nidulans; different antisera, the crossreacting strains can be divided into Coch, Cochliobolus maydis; cra, N. crassa; D, Dodge; dod, N. dodgei; three groups, with members of each group exhibiting reactions gal, N. galapagosensis; Gel, Gelasinospora cerealis; Hon, Honduras; fusion) with one another and lin, N. lineolata Pod, Podospora anserina; Sac, Saccharomyces of complete identity (end-to-end cerevisiae; sit, N. sitophila; Sor, Sordaria fimicola; terri, N. terricola; reactions of only partial identity (indicated by spur formation) tet, N. tetrasperma; tor, N. toroi; W, Warcup. with members of other groups. Thus, N. crassa, N. crassa (Honduras), N. sitophila, N. intermedia, N. tetrasperma (Dodge and Warcup), and N. toroi fall in one group; N. afri- eluate was dialyzed against the same buffer, and tested for cana, N. dodgei, N. galapagosensis, N. fineolata, N. terricola, electrophoretic purity by polyacrylamide gel electrophoresis. and Gelasinospora cerealis in another; and Sordariafimicola Antisera against the antigen were produced in New Zealand falls in still another group. White rabbits by subcutaneous and intramuscular injection of Polyacrylamide gel electrophoresis of perithecial extracts of a 1:1 mixture of antigen and Freund's complete adjuvant the various Neurospora species and Gelasinospora cerealhs (Difco). reveals a major proteinaceous band in each case, with a number Two injection protocols were followed: (i) The rabbits were of electrophoretic variants of the protein occurring (Fig. 2). The injected weekly for 4 weeks and bled 5 weeks after the start of correlation of this major protein with the crossreacting antigen the injections. (ii) Two weekly injections were done, and sera for each of the strains tested was demonstrated by immuno- were collected 6 weeks after the first injection. Identical ex- logical analysis of gel slices taken along the length of unstained perimental results were obtained with sera produced by either gels, inasmuch as only slices corresponding to the major band protocol. To the sera, sodium azide to a final concentration of reacted with the immune sera. Polyacrylamide gels of peri- 0.1% (wt/vol) was added as a preservative, and the sera were
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