68 Worsdell. — The Struchtre and Morphology of the ‘ Ovule ! 
Caspary (25), in a paper, accompanied by beautiful illustrations, on 
abnormal ovules of Trifolium repens, states that the funicle in this plant is 
morphologically equivalent to the lower part of the leafy lobe or pinna of 
the carpel ; at no stage is any limit between the funicle and the outer 
integument to be found. The inner integument, being so similar, must 
possess the same meaning and origin. It may disappear in the tissue 
of the leaflet, just as the outer integument has done in the funicle. ‘The 
funicle, along with the integuments, is in Trifolium repens the morphological 
equivalent of a leaflet (pinna), whose stalk or midrib is in the lower part 
of the funicle and whose campanulate or conical outgrowths of the upper part 
are the integuments. The nucellus is the new shoot seated on this leaflet.’ 
He thinks the nucellus also probably forms an integral part of the carpel. 
Through that classical work of his, the ‘ Bildungsabweichungen,’ 
Cramer (30) established his reputation as one of the most distinguished 
exponents of the morphological nature of the ovule. He observed meta- 
morphosed ovules of Primulaceae ; these were in the form of small foliar 
organs, developed on the free central placenta, which he regarded as entire 
leaves. He regards the nucellus as the product of a leaf. As for the 
integuments, if, he says, they were two distinct leaves and not part of one 
leaf they should both proliferate. That they are not such is shown by 
their becoming concave towards the upper side. The metamorphosed 
ovules are to be regarded as ‘ the outcome of the working of ideal successive 
combinations of formative forces fighting against each other, and not as 
developmental stages which have succeeded each other from time to time.’ 
He is strongly in favour of the use of abnormalities for the solution of 
morphological problems. To him, the ovule of Compositae is, like that of 
Primulaceae, an entire leaf. He also observed proliferous flowers of 
Delphinium elatum in which the green carpels bore either sterile or vegeta- 
tively developed ovules in the form of lobes ; the latter he held to be 
integumental in nature and the protuberances borne by them nucelli. In 
his paper ‘ On the Morphological Significance of the Ovule,’ occurring in 
the same volume, he sets forth that the nucellus is a new structure and not 
of the nature of a shoot, as he does not believe that the latter, as asserted, 
could ever arise from an ovule. The nucellus is merely an emergence , like 
pollen-sacs and fern-sporangia. His researches in the development of the 
ovule in Centaur ea jacea , Lysimachia punctata , and Anther icum liliago 
show that the primary papilla is not the nucellus but later gives rise to both 
this and the integuments. It is the funicle which first appears, and on this 
the nucellus is developed laterally. From his observations on Trifolium 
repens he concludes, with Caspary, that it is the outer, and not the inner 
integument which proliferates and which bears the nucellus in the middle 
of its upper surface. 
Prantl (53) stands conspicuous as one of the bold, yet not necessarily 
