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The Occurrence and Function of Oidia in the Hymenomycetes 1

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The Occurrence and Function of Oidia in the Hymenomycetes 1 THE OCCURRENCE AND FUNCTION OF OIDIA IN THE HYMENOMYCETES 1 Harold J. Brodie IN THE latter half of the eighteenth century the radiatus. He believed that the spermatia became early mycologists, influenced by their knowledge of attached to and emptied their contents into the vesic­ higher plants, searched for sexual organs in the fungi ular female cells. The female cells, after being fertil­ analogous to those with which they were familiar ized, gave rise to branched hyphae which developed among the phanerogams. Micheli (1729) had declared into the fruit body. The observations of contem­ that the cystidia in the hymenium of a mushroom porary workers, such as Eidam (1875) and Kirchner are apetalous flowers. Bulliard (1791) believed the (1875), seemed to support the report of Van Tieghem. cvstidia to be male organs and described the manner However, Van Tieghem (1876) was obliged to in which their seminal fluid poured out over the change his views. He sowed the spermatia of Copri­ basidia to fertilize the basidiospores. nus plicatilis in a drop of nutrient medium and saw It has long been known that the mycelium of many the small cells germinate, producing branched mycelia. fungi breaks up into short segments generally called Because the small cells germinated, he declared that oidia. Such commonly occurring structures as oidia they do not serve as spermatia but reproduce the did not pass unnoticed in the search for sexual organs fungus in an asexual manner, in this respect being and the questions of the significance and probable comparable with conidia. The fact that the oidia did function of oidia engaged the attention of a number sometimes fuse with large vesicular cells was of no of workers. Tulasne (1851) discovered in the lichens special significance, Van Tieghem decided, since the and in many of the Ascomycetes structures in which oidia could fuse equally well with any cell of the were borne small spores usually incapable of germina­ mycelium. He concluded that the supposed sexual tion. These spores he called spermatia and he con­ organs in the Basidiomycetes are in reality merely sidered that they are male cells. Later he discovered vegetative structures. similar structures in the Tremellales and concluded Cornu (1875) succeeded in germinating the so­ that the pycniospores of the Uredinales are also true called spermatia of certain of the Ascomycetes and male cells (Tulasne, 1859). he, like Van Tieghem, considered that the spermatia Hofmeister (1852) showed that in ferns the sex serve only for vegetative reproduction and are com­ organs are developed not on the spore-bearing genera­ parable with conidia. tion but on the prothallium which is independent of This idea was championed by Brefeld (1877, 1889), the sporophyte. This discovery provided mycologists who discredited theories of sexuality in the Basidio­ with a new clue and they began to search for sexual mycetes and who saw no difference between the origin organs in the higher fungi not on the pileus but on of basidiospores and that of conidia. His attention the vegetative mycelium. was much attracted by oidia and he demonstrated Karsten (1860) described finding on the mycelium their manner of occurrence in more than thirty species of Psalliota campestris reniform" female" cells which of the Hymenomyeetes." He announced (1877) that were fertilized by "male" elements. At the same although he had made numerous attempts to do so, time Oersted (1865) thought he saw female cells he had been unable to germinate the oidia of Coprinus fertilized by ordinary hyphae. The suggestion that the commonly occurring oidia lagopus, and from this he concluded that oidia are might be male cells was made again by Rees (1875), vestigial structures which no longer possess the power who germinated basidiospores of Coprinus stereo­ of germinating and which, therefore, cannot be re­ garded as functional spermatia. rarius 2 and observed that, on the young mycelia, there were developed short branches bearing small Richard Falck (1902), a student of Brefeld, demon­ cells which became detached from the parent fila­ strated that oidia are to be found on the mycelium ment. These cells failed to germinate and Rees be­ of species of Mucor, Dacryomyces, Ascobolus, Copri­ lieved that they must be spermatia. Seeing large nus, Collybia and Hypholoma. Like Brefeld, Falck vesicular cells on the same mycelium, he thought he concluded that the oidia of most of the coprophilous had also found the female organs. species of the Basidiomycetes have lost the ability to Van Tiezhem (1875a. b) reported similar spermatia germinate. and female organs in Coprinus ephemeroides and C. The discovery of Schmitz (1880) of paired nuclei in the mycelium and urediniospores of Coleosporium 1 Received for publication February 14. 1935. Papers from the Department of Botany and the campanulae was the begininng of a long series of Herbarium of the University of Michigan, No. 526. cytological investigations which threw light upon the 2 Both Rees and Van Tieghem used the name Coprinus question of sexuality in the Basidiomycetes. Through stercorarius to apply to an oidium-producing funzus, the researches of Rosen, Dangeard, Sappin-Trouffy, Brefeld believed that these workers probably had refer­ Xlairo, and others there evolved a new theory of ence to C. laqopu», The truo C. stercorarius does not produce oidia and is homothallic, " For a list of these species, see Brodie, H. J. (1931). [The JOURXAL for April (23: 235-308) was issued June 15, 1936,] A~mRICAN JOCR:"AL OF BOTANY, VOL. 2:3, xo. 3, )1.", 1936 309 310 AMEIUCAN JOURNAL OF BOTANY [Vo\. 23, sexuality which was announced by Bensaude (1918) ject. However, the researches referred to below will and independently by Kniep (1917). suffice to show how the more recent developments in By cytological and experimental investigations, experimental work concerning sexuality in the higher Bensaude showed that Coprinus lagopus 4 is hetero­ fungi are connected with the problem of the function thallic. She obtained two cultures, each of which was of the oidia. derived from a single basidiospore. These mycelia Vandendries (923) published an account of the remained for eight months in the primary or haploid sexual behavior of the heterothallic agarics Collybia condition-i.e., they produced no clamp-connections, velutipes, Hypholoma fasciculare, and Panaeolus cam­ had but a single nucleus in each cell, and produced panulatus and showed that oidia are produced on the no fruit bodies whatever. However, when the two monocaryon mycelium of each of these species. Some mycelia were allowed to fuse, there was developed a oidia of P. campanulatus were isolated from a mycel­ secondary or dicaryon mycelium 5 which bore clamp­ ium called Thallus 17, a strain which could be recog­ connections. Bensaude showed that the paired nuclei nized by certain specific cultural characters. When of the dicaryophyte divide conjugately and that each the oidia taken from this mycelium were sown on division of the conjugate pair or dicaryon is associated fresh culture medium, a mycelium was obtained which with the formation of a clamp-connection. The two· bore the characters of the parent. From this and nuclei which fuse in the basidium are the descendants other experiments, Vandendries concluded: "L'oidie, of a single pair of nuclei belonging to compatible element haploide, peut jouer un role de multiplica­ strains. tion purement vegetative et propager un thalle pri­ Bensaude also studied the structure and develop­ maire gardant les proprietees de son ascendant: elle ment of the oidia of Coprinus lagopus. In certain peut aussi fertiliser un thalle haploide de sexe con­ culture tubes she saw some of the oidia germinate traire, et presente alars tous les attributs d'un gamete." and produce short germ tubes which exhibited the Craigie (1927) discovered that the pycniospores of ability to fuse with any of the cells of the parent the Uredinales are functional. He found that the mycelium. She had also noticed that when she pycniospores are effective diploidizing agents after planted two mycelia of opposite strain, A and B, near transference by hand or by insects from a (+) my­ one another on nutrient agar, occasionally one of the celium to a (-) mycelium, or vice versa. mycelia became binucleate before it had come into Professor A. H. R. Buller suggested to the writer contact with the other one. She explained this occur­ that the oidia of the Hymenomycetes, like the pycnio­ rence by supposing that the oidia from one mycelium, spores of the Uredinales, after transference from a B, floated across the gap between A and B in the mycelium of one strain to a mycelium of compatible surface film of water covering the agar. When the strain might germinate, and the germ tubes or mycelia germ tubes of the oidia from mvcelium B fused with which developed from the oidia might bring about mycelium A, the latter was co~verted into dicaryon the diploidization of the mycelium to which they had mycelium. been transferred. This suggestion was verified by the As a result of her investigations, Bensaude declared writer using the well known agaric Coprinus lagopus that the oidia of Coprinus lagopus may be the means (Brodie, 1931). whereby the monocaryon mycelium is transformed Later, Dowding (931) found that in one of the into dicaryon mycelium. Ascomycetes, Ascobolus stercorarius, the oidia func­ Since 1918 much attention has been given to the tion in a way similar to those of the Hymenomycetes. study of sexuality in the Hvmenomvcetes, It is not An interesting extension of this idea is the research the writer's intention to review in this paper all the of Drayton (1932) on Sclerotinia Gladioli Massey. recent literature dealing with this very complex sub- It was shown that when this fungus is isolated from Gladiolus and other cormous and bulbous plants, my­ 4 Bensaude referred to this fungus as C..fimeiarius, which has been shown to be identical with the C.
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