516 FILICALES 



round numbers 60, spore-mother-cells traversed. Tangential sections sho\v, 

 on the average, about six such layers of cells to be present, and the 

 average number of spore-mother-cells in each sporangium would thus be 

 6x60 = 360, while the average potential output of spores per sporangium 

 would be 360x4=1440 spores. It may be noted that, in the cases 

 observed, all the spore-mother-cells undergo the tetrad-division, and none 

 have been seen to be disorganised, though this, doubtless, may occur 

 occasionally. 



Reverting to the tissues outside the sporogenous group, the tapetum is 

 not a strictly defined layer, and is often irregularly doubled by periclinal 

 divisions ; but this is by no means always the case. It remains recognis- 

 able as distinct, enlarged, glandular cells, often with several nuclei, up to 

 the period of the tetrad-division. The wall of the sporangium outside the 

 tapetum consists of two or more layers, commonly three, of which the 

 outermost is the firmest and most differentiated : the inner layers are less 

 regular, and are composed of more or less tabular cells. I have never 

 seen the tapetum in direct juxtaposition with the superficial layer, as it 

 has been figured and described by Campbell. 1 Of the external wall three 

 essential parts may be recognised, and they will be best seen in Fig. 284 D, 

 which represents the apex of the sporangium, cut off in such a plane 

 as x, x in Fig. 284 E. The first part (a, Fig. 284 E) is found on the 

 peripheral side of the sporangium, and consists of large turgid cells, 

 with moderately thin walls, and granular contents, which stain deeply 

 with Bismarck brown. A second region (/>) consists of deep prismatic 

 cells with thick lignified walls, which may be recognised as the annulus ; 

 it extends as a narrow bridge across the apex of the sporangium (Fig. 284 D), 

 and widens out on either side, as the apex is left, into a broader band 

 of cells with lignified walls (Fig. 284 F). The third region (c] consists 

 of thinner-walled cells, of elongated form, which constitute a narrow 

 band running down the anterior (ventral) side of the sporangium. This 

 is the tissue which defines the fissure of dehiscence. 



The structure of the sporangial wall being as thus described, we 

 may now consider how it works in connection with dehiscence. The 

 annulus, together with the two broader lateral extensions of it, constitute 

 a firm resistant arch, of which the apex is the narrowest part, being 

 only about three cells wide (Figs. 284 D, E). If the thinner-walled posterior 

 region (a) were to contract, as we may presume it does by drying as the 

 sporangium matures, the two sides would be pulled backwards, while the 

 thin bridge of the annulus at the apex would act as a sort of semi-rigid 

 hinge ; the line of dehiscence on the ventral face, having been structurally 

 defined, would thus, on fission, be caused to gape widely. It is not 

 probable, however, that this hinge-like action is very considerable, and 

 the gaping of the slit may be mainly due, as in other Marattiaceae, to 

 mere drying up of the cells in the neighbourhood of the rupture. 



iS and /< r/tx, p. 297, and Fig. 1640. 



