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									The Angiosperm Seed
Develops from the ovule as a result of double fertilisation. Contains the embryo and stored
nutrients, and is protected by the seed coat (testa). The seed coat is derived from the
Endosperm: derived from the fused polar and sperm nuclei in the central cell of the embryo
sac, stores food external to the embryo.
Perisperm: sporophyte nucellar tissue, stores food external to the embryo.
In many dicots, the endosperm and perisperm are transient and are absorbed by the
developing embryo before the seed becomes dormant. The mature seed then stores its
food in the cotyledons which become fleshy.
The micropyle may remain as an occluded pore or disappear. The funiculus abscises from
the ovule, leaving a scar (hilum). In anatropous ovules part of the funiculus remains as a
longitudinal ridge, the raphe.
Aril: a funiculus outgrowth. Caruncle: an integumentary protuberance near the micropyle.
Elaiosome: an oily appendage used as food by ants.

Seed Development
Double fertilisation initialises seed development. The ovule and endosperm grow and the
embryo sac enlarges to accommodate the endosperm. Once the endosperm reaches
maximum volume, the embryo starts to grow rapidly, initially by cell division and then by cell
enlargement. In the pea (Pisum sativum), the embryo almost fills the embryo sac as it
grows at the expense of the endoderm. (Organic C is tranferred from the endosperm to the
embryo). Finally the seed becomes dormant (at this stage in the pea the suspensor
The growing embryo sac is a sink, absorbing water and soluble materials from the
surrounding ovular tissues (which are digested) which are supplied by the vessels in the
funiculus. In the pea, the growth of pod and seeds is under the control of gibberellins, auxin
and abscisic acid. Hormones mobilised synthesised by the growing ovules, following
fertilisation, stimulate growth of the pod.
Albuminous seeds: store food in the endosperm (monocotyledons) or perisperm
(Amaranthaceae, Chenopodiaceae, Polygonaceae).
Exalbuminous seeds: food is stored in the embryo; no endosperm or perisperm when
Most seeds show a combination of storage tissue types.

Mature Embryo
Dicotyledons: the two cotyledons arise as lateral organs at the apex of the embryo axis and
are in the same position, relative to the meristem, as foliage leaves.
Monocotyledons: one cotyledon, displaces apical meristem to a lateral position.
Embryo: upright, bent or curved.
                           Breaking Seed Dormancy

                                                                       Seed coat (testa)

 Cotyledon (scutellum)
                                                                         Aleurone layer
                                                       amylase           Storage protein

                                                       RNA                 Amino acids

                                                   DNA                        660 nm
                                                                        Pfr             Pr
            Radicle                                                           730 nm

                  H2O                                                           Dark


The diagram above shows the breaking of seed dormancy (in a generalised grass/cereal
All seeds must absorb (imbibe) water to become metabolically active, but not all require a
red light stimulus to stimulate phytochrome, e.g. lettuce seeds require both the red light
stimulus and gibberellic acid activation by imbibition, whereas barley seeds require only
gibberellic acid activation by imbibition.
Q.1 At what two stages on the above diagram is hydrolysis occurring?
Q.2 What is the function of the scutellum in grass seeds?
Q.3 What is the advantage of requiring a red light stimulus to break dormancy in certain
Some plants require a cold period to break seed dormancy, and others require damage to
the seed coat to allow water to enter. In the walnut (Juglans regia) the nut has to
decompose on the forest floor for two years, before its hard seed coat is weakened enough
to allow water in and the germinating embryo out!
Endosperm Formation
Endosperm is characteristic of angiosperm seeds. There are three modes of endosperm
1. Nuclear: many nuclei are formed by nuclear divisions whilst the endosperm remains
noncellular or develops cell walls later. The centre of the cell is occupied by a large vacuole
and the nuclei are parietal. In Capsella, endosperm nuclei accumulate at the two ends of
the embryo sac and the endosperm at the chalazal end digests nucellar cells in front of the
advancing embryo sac. The nucellar cells hypertrophy before digestion and become rich in
protein and nucleic acids. Their digested remains are seen inside vesicles in large vacuoles
in the endosperm cells. In Eranthis hiemalis (Ranunculaceae) the endosperm digests
adjacent integumentary cells. Integumentary cell nuclei migrate into the endosperm and are
broken down. This reduces the integumentary cell layers from 11 to 7 by the time the
endodermis becomes cellular. The cell walls form from phragmoplasts or by wall ingrowth.
In Pisum, an endosperm cell wall and middle lamella joins the endosperm from the outer
walls of the embryo. The outermost layer becomes the aleurone layer and endosperm cells
eventually take up the central vacuolar space. In Cocos nucifera, the central cavity does
not fill with cells, but contains coconut milk.
2. Cellular: cell wall formation begins with the first mitosis and continues until the
endosperm stops growing.
3. Helobial: the embryo sac is divided into two unequal cells, the larger chalazal cell usually
develops noncellularly whilst the smaller micropylar cell develops in either manner. Occurs
mainly in the monocotyledons.
Food stores contain mainly carbohydrate, protein and lipid. Cereals: 70-80% dry weight is
starch, peas and beans 50% starch. Zea contains mostly starch in the endosperm, but the
embryo is 50% oil. Rape and mustard (Brassicaceae) are 40% oil and 30% protein; soya
(Fabaceae) seeds are 20% oil and 40% protein. The starchy endosperm may be living, with
starch grains inside amyloplasts, or nonliving with free starch grains. Carbohydrates may
also be stored in thickened cell walls (of endosperm or cotyledons) composed mostly of
hemicelluloses. Protein is stored in granules enclosed in membrane derived from the
tonoplast and may consist of globulins. Oil is stored as triglyceride in cytoplasmic granules,
which may be bound by a unit membrane or a phospholipid monolayer (?).

Seed Coat
The seed coat is dry in most angiosperms, but may have fleshy appendages, e.g.
elaiosomes, or juicy layers such as the fleshy epidermis of pomegranate (Punica) seeds.
Many gymnosperms have fleshy seed coats. Cuticular layers and phenolic compounds
restrict water entry into the seed.
The seed coat and associated coverings (pericarp, other floret parts in grasses) may
contain germination inhibitors and only when these are removed will the seed germinate.
Seeds that rely on animal ingestion to disperse them, have testa that are resitant to
digestive processes. The epidermis of some seeds secretes mucilage and this may cause
them to stick to animals and be dispersed, or may prevent desiccation or prevent
germination in excessive moisture by swelling and impeding oxygen diffusion into the seed.

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