ANGIOSPERMS. 
573 
the last remaining doubt on this points Cramer has shown in a number of 
other instances that all stages of the metamorphosis of ovules occur when the 
flower is developed in a monstrous condition, leading also to the conclusion that 
the nucellus is a lateral development on the funiculus of the ovule. Malformations 
of Delphinium elatum, where the ovules spring from the margins of the carpels, show 
that the carpel is transformed into a flat open pinnate leaf, the lobes of which are 
the metamorphosed ovules. The nucellus here springs from the upper or inner 
side of the lobe of the leaf which represents the transformed funiculus together 
with the integument. In Melilotus, Primula chinaisis, and Umbelliferae, Cramer 
found the same to be the case ^ Relying on this and other facts, and on the 
hypothesis that the ovule is never a terminal structure of the floral axis, Cramer-^ 
adopted the view that the ovule is either a metamorphosed leaf or part of a leaf 
(a tooth or outgrowth of the upper surface). The ovule of Primulaceae and Com- 
positae he considered to be a whole leaf, and he supposed that closer observation 
would show the same to be the case in other flowers also, especially in those 
where the flower is said to possess a solitary ' reputed terminal ovule,' as Urtica 
(and Taxus), and perhaps also the Dipsacaceaä and others. The nucellus would in 
this case be a new formation on the surface of the ovular leaf, the funiculus would 
correspond to the base of this leaf, and the integuments to its upper part, which is 
folded once or twice in the form of a cup or hood round the nucellus. On the 
other hand he would consider as only portions of the leaf (teeth or outgrowths of 
the upper surface) all those ovules which spring singly or in numbers from the 
margin or upper surface of carpellary leaves, as those of Cycadeas, Abietinese (?), 
Liliaceae, Umbelliferae, Ranunculaceae, Resedaceae, Cruciferae, Leguminosae, &c. In 
these cases the nucellus would be a new formation on the surface of the lobe, the 
funiculus would correspond to its base, and the integuments to its upper part folded 
once or twice round the nucellus in the form of a cup. Only in those few plants in 
which the ovule has no integument would the naked nucellus or entire ovule corre- 
spond to this lobe of the carpellary leaf. In the first edition of this book I expressed 
my agreement with Cramer's view, but with a reservation with respect to Orchideae, 
being especially influenced by the importance which I then attached to the morpho- 
logical equivalency of the nucellus in all Phanerogams. Further reflection has, 
however, deprived this reason of its importance; and I am the more induced to 
ascribe different morphological significations to the ovules, according to their mode 
of origin and their position, because (as has been shown by Magnus, Rohrbach, 
Hanstein, and Schmitz *) in Piperacese, Typhaceae, and Naiadeae the ovule is 
actually the terminal structure of the floral axis, and in Naias this terminal ovule 
is also anatropous. In these statements I not only find the confirmation of my 
^ Schenk writes, ' that which appears plausible enough in the Composite is certainly not the 
case in other families ; the ovule is not a lateral branch of the primitive rudiment, but this itself 
developes into the ovule.' 
2 Compare also H. von Mohl, Vermischte Schriften, pi. I. figs, 27-29. 
^ Cramer, Bildungsabweichungen bei einigen wichtigeren Pflanzenfamilien und die morpho- 
logische Bedeutung des Pflanzeneies (Zürich 1869, p. 120), where the literature of this subject has 
been carefully treated, 
* These researches have been already quoted; [see also Eichler, Helosideen, Bot. Zeitg. 1868, 
and Strasburger, Coniferen und Gnetaceen, 1872.] 
