Fig. 410.
Chromosomes uniting at
poles to form the nuclei
of the four spores.
(After Mottier.)

The number of chromosomes usually the same in a given species throughout one phase of the plant.—In those plants which have been carefully studied, the number of chromosomes in the dividing nucleus has been found to be fairly constant in a given species, through all the divisions in that stage or phase of the plant, especially in the embryonic, or young growing parts. For example, in the prothallium, or gametophyte, of certain ferns, as osmunda, the number of chromosomes in the dividing nucleus is always twelve. So in the development of the pollen of lilium from the mother cells, and in the divisions of the antherid cell to form the generative cells or sperm cells, there are always twelve chromosomes so far as has been found. In the development of the egg of lilium from the macrospore there are also twelve chromosomes.

When fertilization takes place the number of chromosomes is doubled in the embryo.—In the spermatozoid of osmunda then, as well as in the egg, since these are developed on the gametophyte, there are twelve chromosomes each. The same is true in the sperm cell (generative cell) of lilium, and also in the egg-cell. When these nuclei unite, as they do in fertilization, the paternal nucleus with the maternal nucleus, the number of chromosomes in the fertilized egg, if we take lilium as an example, is twenty-four instead of twelve; the number is doubled. The fertilized egg is the beginning of the sporophyte, as we have seen. Curiously throughout all the divisions of the nucleus in the embryonic tissues of the sporophyte, so far as has been determined, up to the formation of the mother cells of the spores, the number of chromosomes is usually the same.

Fig. 411.

Karyokinesis in sporophyte cells of podophyllum (twice the number of chromosomes here that are found in the dividing spore mother cells).

682. Reduction of the number of chromosomes in the nucleus.—If there were no reduction in the number of chromosomes at any point in the life cycle of plants, the number would thus become infinitely large. A reduction, however, does take place. This usually occurs, either in the mother cell of the spores or in the divisions of its nucleus, at the time the spores are formed. In the mother cells a sort of pseudo-reduction is effected by the chromatin band separating into one half the usual number of nuclear segments. So that in lilium during the first division of the nucleus of the mother cell the chromatin band divides into twelve segments, instead of twenty-four as it has done throughout the sporophyte stage. So in podophyllum during the first division in the mother cell it separates into eight instead of into sixteen. Whether a qualitative reduction by transverse division of the spirem band, unaccompanied by a longitudinal splitting, takes place during the first or second karyokinesis is still in doubt. Qualitative reduction does take place in some plants according to Beliaieff and others. Recently the author has found that it takes place in Trillium grandiflorum during the second karyokinesis, and in Arisæma triphyllum the chromosomes divide both transversely and longitudinally during the first karyokinesis forming four chromosomes, and a qualitative reduction takes place here.

683. Significance of karyokinesis and reduction.—The precision with which the chromatin substance of the nucleus is divided, when in the spirem stage, and later the halves of the chromosomes are distributed to the daughter nuclei, has led to the belief that this substance bears the hereditary qualities of the organism, and that these qualities are thus transmitted with certainty to the offspring. In reduction not only is the original number of chromosomes restored, it is believed by some that there is also a qualitative reduction of the chromatin, i.e. that each of the four spores possesses different qualitative elements of the chromatin as a result of the reducing division of the nucleus during their formation.

The increase in number of chromosomes in the nucleus occurs with the beginning of the sporophyte, and the numerical reduction occurs at the beginning of the gametophyte stage. The full import of karyokinesis and reduction is perhaps not yet known, but there is little doubt that a profound significance is to be attached to these interesting phenomena in plant life.