Active growth in the embryo, other than swelling resulting from imbibition, usually begins with the emergence of the primary root, known as the radicle, from the seed, although in some species (e.g., the coconut) the shoot, or plumule, emerges first. Early growth is dependent mainly upon cell expansion, but within a short time cell division begins in the radicle and young shoot, and thereafter growth and further organ formation (organogenesis) are based upon the usual combination of increase in cell number and enlargement of individual cells.
Until it becomes nutritionally self-supporting, the seedling depends upon reserves provided by the parent sporophyte. In angiosperms these reserves are found in the endosperm, in residual tissues of the ovule, or in the body of the embryo, usually in the cotyledons. In gymnosperms food materials are contained mainly in the female gametophyte.
Since reserve materials are partly in insoluble form—as starch grains, protein granules, lipid droplets, and the like—much of the early metabolism of the seedling is concerned with mobilizing these materials and delivering, or translocating, the products to active areas. Reserves outside the embryo are digested by enzymes secreted by the embryo and, in some instances, also by special cells of the endosperm.
In some seeds (e.g., castor beans) absorption of nutrients from reserves is through the cotyledons, which later expand in the light to become the first organs active in photosynthesis. When the reserves are stored in the cotyledons themselves, these organs may shrink after germination and die or develop chlorophyll and become photosynthetic.
Environmental factors play an important part not only in determining the orientation of the seedling during its establishment as a rooted plant but also in controlling some aspects of its development. The response of the seedling to gravity is important. The radicle, which normally grows downward into the soil, is said to be positively geotropic.
The young shoot, or plumule, is said to be negatively geotropic because it moves away from the soil; it rises by the extension of either the hypocotyl, the region between the radicle and the cotyledons, or the epicotyl, the segment above the level of the cotyledons. If the hypocotyl is extended, the cotyledons are carried out of the soil. If the epicotyl elongates, the cotyledons remain in the soil.
Light affects both the orientation of the seedling and its form. When a seed germinates below the soil surface, the plumule may emerge bent over, thus protecting its delicate tip, only to straighten out when exposed to light (the curvature is retained if the shoot emerges into darkness). Correspondingly, the young leaves of the plumule in such plants as the bean do not expand and become green except after exposure to light. These adaptative responses are known to be governed by reactions in which the light-sensitive pigment phytochrome plays a part.
In most seedlings, the shoot shows a strong attraction to light, or a positive phototropism, which is most evident when the source of light is from one direction. Combined with the response to gravity, this positive phototropism maximizes the likelihood that the aerial parts of the plant will reach the environment most favourable for photosynthesis.