Chapter 36), and at the outer end of the raphe is the _chalaza_,
the point where the stalk is joined to the end of the ovule, best understood in a straight ovule. Upon the opposite side of the scar and close to it can be seen a minute depression, the _micropyle_. Underneath the seed coat and lying between this point and the end of the seed is the _embryo_, which gives greater prominence to the bean at this point, but it is especially more prominent after the bean has been soaked in water. Soak the beans in water and as they are swelling note how the seed coats swell faster than the inner portion of the seed, which causes them to wrinkle in a curious way, but finally the inner portion swells and fills the seed coat out smooth again. Sketch a bean showing all the external features both in side view and in front. Split one lengthwise and sketch the half to which the embryo clings, noting the young root, stem, and the small leaves which were lying between the cotyledons. There is no endosperm here now, since it was all used up in the growth of the embryo, and a large part of its substance was stored up in the cotyledons. As the seed germinates the young plant gets its first food from that stored in the cotyledons. The hypocotyl elongates, becomes strongly arched, and at last straightens up, lifting the cotyledons from the soil. As the cotyledons become exposed to the light they assume a green color. Some of the stored food in them goes to nourish the embryo during germination, and they therefore become smaller, shrivel somewhat, and at last fall off.
=211. The castor-oil bean.=—This is not a true bean, since it belongs to a very different family of plants (Euphorbiaceæ). In the germination of this seed a very interesting comparison can be made with that of the garden bean. As the “bean” swells the very hard outer coat generally breaks open at the free end and slips off at the stem end. The next coat within, which is also hard and shining black, splits open at the opposite end, that is at the stem end. It usually splits open in the form of three ribs. Next within the inner coat is a very thin, whitish film (the remains of the nucellus, and corresponding to the perisperm) which shrivels up and loosens from the white mass, the endosperm, within. In the castor-oil bean, then, the endosperm is not all absorbed by the embryo during the formation of the seed. As the plant becomes older we should note that the fleshy endosperm becomes thinner and thinner, and at last there is nothing but a thin, whitish film covering the green faces of the cotyledons. The endosperm has been gradually absorbed by the germinating plant through its cotyledons and used for food.
Arisæma triphyllum.[15]
=212. Germination of seeds of jack-in-the-pulpit.=—The ovaries of jack-in-the-pulpit form large, bright red berries with a soft pulp enclosing one to several large seeds. The seeds are oval in form. Their germination is interesting, and illustrates one type of germination of seeds common among monocotyledonous plants. If the seeds are covered with sand, and kept in a moist place, they will germinate readily.
=213. How the embryo backs out of the seed.=—The embryo lies within the mass of the endosperm; the root end, near the smaller end of the seed. The club-shaped cotyledon lies near the middle of the seed, surrounded firmly on all sides by the endosperm. The stalk, or petiole, of the cotyledon, like the lower part of the petiole of the leaves, is a hollow cylinder, and contains the younger leaves, and the growing end of the stem or bud. When germination begins, the stalk, or petiole, of the cotyledon elongates. This pushes the root end of the embryo out at the small end of the seed. The free end of the embryo now enlarges somewhat, as seen in the figures, and becomes the bulb, or corm, of the young plant. At first no roots are visible, but in a short time one, two, or more roots appear on the enlarged end.
=214. Section of an embryo.=—If we make a longisection of the embryo and seed at this time we can see how the club-shaped cotyledon is closely surrounded by the endosperm. Through the cotyledon, then, the nourishment from the endosperm is readily passed over to the growing embryo. In the hollow part of the petiole near the bulb can be seen the first leaf.
=215. How the first leaf appears.=—As the embryo backs out of the seed, it turns downward into the soil, unless the seed is so lying that it pushes straight downward. On the upper side of the arch thus formed, in the petiole of the cotyledon, a slit appears, and through this opening the first leaf arches its way out. The loop of the petiole comes out first, and the leaf later, as shown in fig. 98. The petiole now gradually straightens up, and as it elongates the leaf expands.
=216. The first leaf of the jack-in-the-pulpit is a simple one.=—The first leaf of the embryo jack-in-the-pulpit is very different in form from the leaves which we are accustomed to see on mature plants. If we did not know that it came from the seed of this plant we would not recognize it. It is simple, that is it consists of one lamina or blade, and not of three leaflets as in the compound leaf of the mature plant. The simple leaf is ovate and with a broad heart-shaped base. The jack-in-the-pulpit, then, as trillium, and some other monocotyledonous plants which have compound leaves on the mature plants, have simple leaves during embryonic development. The ancestral monocotyledons are supposed to have had simple leaves. Thus there is in the embryonic development of the jack-in-the-pulpit, and others with compound leaves, a sort of recapitulation of the evolutionary history of the leaf in these forms.
=216=_a_. =Germination of the pea.=—Compare with the bean. Note especially that the cotyledons are not lifted above the soil as in the beans. Compare germination of acorns.
Digestion.
=216=_b_. =To test for stored food substance in the seedlings studied.=—The pumpkin, squash, and castor-oil bean are examples of what are called oily seeds, since considerable oil is stored up in the protoplasm in the cotyledons. To test for this, remove a small portion of the substance from the cotyledon of the squash and crush it on a glass slip in a drop or two of osmic acid.[16] Put on a cover glass and examine with a microscope. The black amorphous matter shows the presence of oil in the protoplasm. The small bodies which are stained yellow are _aleurone_ grains, a form of protein or albuminous substance. Both the oil and the protein substance are used by the seedling during germination. The oil is converted into an available food form by the action of an enzyme called _lipase_, which splits up the fatty oil into glucose and other substances. Lipase has been found in the endosperm of the castor-oil, cocoanut, and in the cotyledons of the pumpkin, as well as in other seeds containing oil as a stored product. The aleurone is made available by an enzyme of the nature of trypsin. Test the endosperm of the castor-oil bean in the same way. Make another test of both the squash and castor-oil seeds with iodine to show that starch is not present.
Test the cotyledon of the bean with iodine for the presence of starch. If the endosperm of corn seed has not been tested do so now with iodine. The endosperm consists largely of starch. The starch is converted to glucose by a diastatic “ferment” formed by the seedling as it germinates. Make a thin cross-section of a grain of wheat, including the seed coat and a portion of the interior, treat with iodine and mount for microscopic examination. Note the abundance of starch in the internal portion of endosperm. Note a layer of cells on the outside of the starch portions filled with small bodies which stain yellow. These are aleurone grains. The cellulose in the cell walls of the endosperm is dissolved by another enzyme called _cytase_, and some plants store up cellulose for food. For example, in the endosperm of the _date_ the cell walls are very much thickened and pitted. The cell walls consist of reserve cellulose and the seedling makes use of it for food during growth.
=216=_c_. =Albuminous and exalbuminous seeds.=—In seeds where the food is stored outside of the embryo they are called _albuminous_; examples, corn, wheat and other cereals, Indian turnip, etc. In those seeds where the food is stored up in the embryo they are called _exalbuminous_; examples, bean, pea, pumpkin, squash, etc.
=217. Digestion= has a well-defined meaning in animal physiology and relates to the conversion of solid food, usually within the stomach, into a soluble form by the action of certain gastric juices, so that the liquid food may be absorbed into the circulatory system. The term is not often applied in plant physiology, since the method of obtaining food is in general fundamentally different in plants and animals. It is usually applied to the process of the conversion of starch into some form of sugar in solution, as glucose, etc. This we have found takes place in the leaf, especially at night, through the action of a diastatic ferment developed more abundantly in darkness. As a result, the starch formed during the day in the leaves is digested at night and converted into sugar, in which form it is transferred to the growing parts to be employed in the making of new tissues, or it is stored for future use; in other cases it unites with certain inorganic substances, absorbed by the roots and raised to the leaf, to form proteids and other organic substances. In tubers, seeds, parts of stems or leaves where starch is stored, it must first be “digested” by the action of some enzyme before it can be used as food by the sprouting tubers or germinating seeds.
For example, starch is converted to a glucose by the action of a diastase. Cellulose is converted to a glucose by cytase. Albuminoids are converted into available food by a tryptic ferment. Fatty oils are converted into glucose and other products by lipase.
Inulin, a carbohydrate closely related to starch, is stored up for food in solution in many composite plants, as in the artichoke, the root tuber of dahlia, etc. When used for food by the growing plant it is converted into glucose by an enzyme, inulase. Make a section of a portion of a dahlia tuber or artichoke and treat with alcohol. The inulin is precipitated into sphæro crystals. (See also paragraphs 156-161 and 216_b_.)
=218.= Then there are certain fungi which feed on starch or other organic substances whether in the host or not, which excrete certain enzymes to dissolve the starch, etc., to bring it into a soluble form before they can absorb it as food. Such a process is a sort of _extracellular digestion_, i.e., the organism excretes the enzyme and digests the solid outside, since it cannot take the food within its cells in the solid form. To a certain degree the higher plants perform also extracellular digestion in the action of root hair excretion on insoluble substances, and in the case of the humus saprophytes. But for them soluble food is largely prepared by the action of acids, etc., in the soil or water, or by the work of fungi and bacteria as described in