The emergence of the seedling

Active growth in the embryo, other than swelling resulting from imbibition, usually
begins with the emergence of the primary root from the seed, although in some species
(e.g., the coconut) the shoot emerges first. Early growth is dependent mainly upon cell
expansion, but, within a short time, cell division begins in the radicle and young shoot;
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, 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, but, 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.
Later development: the sporophyte plant body

Continuation of organ formation

Although it is convenient to refer to the early development of the plant sporophyte from
the fertilized egg as embryogenesis, the process is never actually concluded as it is in the
higher animals. In vascular plants, organ formation (organogenesis) is not confined to
early life, and the processes of shoot, root, and leaf formation that occur first in the
embryo are repeated, albeit in modified form, throughout the life of the plant. The life
span may be short and determinate, as in annual plants such as the cereals, or long,
lasting for many years—indeed potentially indefinitely, except for limitations imposed
by the environment and accidents—as in trees. The protracted growth of perennials, or
plants that resume growth each growing season, tends to lead to increase in size, but
bulk is not necessarily directly correlated with age, because individual leaves, flowers,
and even whole limbs continuously die and are shed. Some long-lived plants, however,
do reach a point at which losses of body mass balance the increase resulting from
continued growth and organ formation.