The Nucellus, Embryo Sac, Endosperm, and Embryo of Aesculus and their Interdependence during Growth

Immature fruits of Aesculus yield powerful stimuli to growthand cell division. Therefore, the developing fruit of Aesculuswoerlitzensis Koehne has been investigated from pollinationto maturity. The fluid, or liquid endosperm, which containsthe growth-promoting substances is produced in a large vesiclewhich forms at the chalazal tip of the embryo sac. As the vesiclegrows, it encroaches upon the nucellus and, when the embryodevelops, one of its cotyledons penetrates into the vesicleof the embryo sac where it grows and absorbs the contents. Theembryo, which has only a vestigial suspensor, reaches the vesicleby growing along the neck of the long curved embryo sac. Thecotyledon which first penetrates the vesicle grows into a massivestructure; the other remains small. The tip of the cotyledonseems to function as an absorbing surface, for the endospermwith which it comes into contact disorganizes. Fertilizationand the presence of a viable embryo at the micropylar end ofthe embryo sac therefore sets in train a number of other events.These are the extensive development of the nucellus at the chalazalend of the embryo sac, the swelling of the vesicle and the formationof a free nuclear and some cellular endosperm, and the disorganizationof the nucellus as it is encroached upon by the vesiculate embryosac. Attention is directed to the organization of the nucellusin the vicinity of the embryo sac. Files or richly protoplasmicnucellar cells(hypostase) which converge upon the chalazal tipof the embryo sac suggest a principal route by which the vesiclemay be nourished. Special attention is drawn to the very differentsizes of cells, their nuclei and nucleoli, in the differentparts of the nucellus. The growth and development of the embryohas also been traced from the zygote to the mature seed. Thenutritive role of the veaiculate embryo sac, and the supplyof growthstimulating substances, through the function of a cotyledonas an absorbing organ, are now seen as important features ofthe development of the Aesulus embryo in the ovule. Many outstandingproblems still remain. The sequential events that follow fertilizationin the different interdependent regions (nucellus, embryo sac,cotyledon, &c.) are here described, but not casually explained.