I 82 DIVISION II. — COURSE OF DEVELOPMENT OF FUNGI. 



is true that conjugation of more than two primary sporidia has been observed in the 

 plants which we are considering ; in Tilletia, for instance, and Ustilago Tragopogonis; 

 but these exceptional cases are comparatively rare and would only confirm the rule, 

 even if similar exceptions in typical sexual conjugation, as in Zygnema and Acctabu- 

 laria, were unknown. 



The reasons which Brcfeld alleges against my conception of the pairing, as far as 

 I understand them, are these. Pairing occurs in germination in water ; it does not 

 occur when the primary sporidia of Tilletia, for example, are placed in nutrient 

 solutions before pairing can begin ; it also does not occur in Ustilago Carbo and 

 similar germinating species, if they germinate and sprout in a nutrient solution. But 

 when the supply of food is exhausted, pairing sets in in many species and is followed 

 by the formation of incipient mycelium. A fresh supply of food again sets up 

 gonidial sprouting. This shows that the uniting cells are not sexual organs in 

 themselves incapable of germination, for ' according to our present ideas with 

 respect to the essence of sexuality ' no such change in the properties of sexual cells 

 could be brought about by changes in the nutrition. The answer to this is, that, 

 according to our present hwivlcdge, the only processes of conjugation which can be 

 here brought into comparison, and which are to be distinctly regarded as simplified 

 analogues of sexual processes, consist solely in the characteristic coalescence in 

 each case of two cells into one capable of further development. What further 

 ' essence ' is involved in the matter we do not know. Many of these conjugating 

 cells are not capable of further development without conjugation, the gametes, for 

 example, of many Chlorophyceae ; but this is not the case with all. I will not 

 mention the Phaeosporeae, which supply examples of the opposite behaviour, because 

 the circumstances in their case are not quite clear and are to some extent open 

 to question. But Rostafinski and Woronin found that the swarm-spores developed 

 from recently matured resting-spores of Botrydium granulatum 1 do typically and 

 necessarily conjugate, and that the same swarm-spores germinate without con- 

 jugation when they proceed from resting-spores which are more than two years 

 old. A change therefore takes place in these cells, affecting those properties in them 

 with which we are at present concerned, through the influence of external causes 

 which must be closely connected with processes of nutrition. Even highly differen- 

 tiated sexual cells may be capable of further development when fertilisation is 

 prevented, as is shown by the oogonia with unfertilised oosphercs of Pythium 

 megalacanthum 2 which send out strong germ-tubes. We see therefore that the 

 exceptional cases to the pairing which is the general rule in the Ustilagineae, whether 

 artificially produced or spontaneous, are not opposed to phenomena known to occur in 

 organs which are undoubtedly sexual. 



That there are species which do not pair cannot affect our decision with regard 

 to those which do; moreover it is well known, that in other groups of Fungi organs 

 strictly homologous have a distinct sexual function in some species and may be 

 asexual in others, as we learn by comparison of the Peronosporeae and those Sapro- 

 legnieae which are without antheridia. 



1 TJot. Ztg. 1S77, p. 662. 

 * Bot. Ztg. 1881, p. 543. 



