Asterad Type.

Lactuca saliva (Jones, 1927) may be used as an illustration of the Asterad type which has been based on the studies of Carano (1915) and Soueges (1920c) on various members of the Compositae. The four-celled proembryo consists of two juxtaposed cells derived from the terminal cell ca and two superposed cells ci and m derived from the basal cell cb (Fig. 148 A-C). In the following stage, each of the four cells divides again so that the terminal tier now comprises the quadrant cells q, the middle tier comprises the two juxtaposed cells at m, and ci divides transversely to form the daughter cells n and n' (Fig. 148D). Thus the upper three tiers of this stage owe their origin to the basal cell cb, and the lowest tier of four cells to the terminal cell ca of the two-celled proembryo. The four cells of the tier q divide to form the octant stage, the walls segmenting the quadrant cells being oriented more or less diagonally ; the two cells of the tier m undergo a vertical division to give rise to four cells lying directly above the octants; n also divides by a vertical wall ; and n' divides by a transverse wall to form o and p ). At the same time tangential walls are laid down in the tiers q and m to cut off an outer layer of dermatogen cells from the inner cells which undergo further divisions to give rise to the periblem and plerome (Fig. 148G). Regarding further development, the cell p gives rise to a suspensor consisting of a variable number of cells; o to the root cap and dermatogen of the root; n to the remaining part of the root tip; m to the hypocotyledonary region; and q to the cotyledons and stem tip. Geum urbanum (Soueges, 19236) offers a significant variation from the above scheme in the early demarcation of a special cell called the epiphysis initial. 5 After the two-celled stage (Fig. 149A) the first wall in ca is markedly oblique, resulting in two unequal cells a and b (Fig. 149B-C). Of these, a divides to cut off a wedge-shaped cell e, which is called the epiphysis (Fig. 149 D,E). At the same time the middle cell m divides vertically and ci divides transversely, so that there are now four tiers of cells in all, designated as q, m, n y and n', lying directly above the epiphysis initial e (Fig. 149F). Of these, q gives rise to the cotyledonary region, m to the hypocotyl,

n or one of its derivatives to the hypophysis and a part of the suspensor, and n' to the greater portion of the suspensor. Some of the stages in development are shown in Fig. 149G-L. An epiphysis initial has also been reported in Fragaria and Viola (Soueges, 19356, 1937c). The former is essentially similar to Geum, but the latter shows some differences in that the derivatives of the tier q, lying just above the epiphysis, produce not only the cotyledonary region but also the lower portion of the hypocotyl, while the tier m contributes to the upper portion of the hypocotyl and the root. A suspensor is virtually absent, as the uppermost tier ci is devoted to the formation of the root cap. Solanad Type. Soueges (19206, 1922) and Bhaduri (1936) have studied a number of species of the Solanaceae, of which Nicotiana may be described here as an example. The first division of the zygote is transverse (Fig. 150 A, B). The terminal cell ca and the basal cell cb both divide transversely to give rise to a four-celled proembryo (Fig. 150C-F). The four tiers may be designated from below upward as I, V , m, and d. Now I and V divide by vertical walls oriented at right angles to each other to give rise to octants, while m and ci divide transversely to produce d, f, n, and n' (Fig. 150 G-H). By subsequent divisions the tier I gives rise to the cotyledonary portion, V to the hypocotyl and to the periblem and plerome of the root, and d to the root tip. The remaining cells/, n, and n' produce the suspensor. Hewitt (1939) has recently given a very full account of the embryogeny of Lobelia amoena, which also belongs to the Solanad type. After the two-celled stage (Fig. 151 A) the basal cell cb divides transversely to produce ci and m (Fig. 1515) and the terminal cell divides to produce I and V (Fig. 151C). Following this four-celled stage, m and ci divide to produce the cells d,f, n, and n' (Fig. 151D), each

of which may divide again, resulting in a suspensor which is about 8 to 12 cells long. At the same time the cell V segments into ph and h (Fig. 151D) after which I and ph divide by longitudinal walls oriented at right angles to each other, and h divides by a transverse wall. Two juxtaposed cells are thus produced from I and ph and two superposed cells (ha and Kb)' from h (Fig. 151E). The embryo proper (excluding the suspensor cells) now consists of four tiers, I, ph, ha, and Kb. In I and ph the next division is vertical and in a plane at right angles to the first division. At the same time ha also divides by a vertical wall, resulting in two tiers of four cells each at I and ph, one tier of two cells at ha, and one of a single cell Kb (Fig. 151F). In the cells of the apical tier /, the next division walls are diagonal, followed by periclinal divisions which cut off the dermatogen. The cells of the tier ph divide periclinally; the outer cells form the dermatogen, and the inner undergo further longitudinal divisions to form the periblem and plerome initials of the stem (Fig. 151 G). The two cells of the tier ha give rise to a single semicircular layer of cells which contributes to the periblem, the dermatogen, and a part of the root cap (Fig. 151H-J). The cell Kb divides only at a comparatively late stage, at first by a transverse wall and then by a vertical wall in each of the daughter cells. The upper derivatives form a part of the root cap, which is supplemented on all sides by a cell from the tier ha and in some cases by extra cells cut off from the dermatogen of tier ph. A peculiarity of the suspensor is the widening and vertical divisions of two or three of its cells lying just above the embryo proper. In Sherardia (Soueges, 1925), a member of the Rubiaceae, the four-celled proembryo arises in the same way as in Nicotiana and Lobelia. The first division in the cells / and V (Fig. 152A) may be either transverse or vertical, but m and ci always divide transversely (Fig. 1525). The eight-celled stage may thus comprise six, seven (Fig. 152D), or eight (Fig. 152C) tiers of cells. Normally the derivatives of I form the cotyledonary region and those of V form the hypocotyl; m and ci give rise to a massive suspensor, which is filamentous towards the apical end but is composed of a number of large vesicular cells at the basal end (Fig. 152E-G). . Presumably the last cell of the suspensor functions as a hypophysis and contributes to a portion of the root tip.