Division and Arrangement of the Experiments.

If two plants which differ constantly in one or several characters be crossed, numerous experiments have demonstrated that the common characters are transmitted unchanged to the hybrids and their progeny; but each pair of differentiating characters, on the other hand, unite in the hybrid to form a new character, which in the progeny of the hybrid is usually variable. The object of the experiment was to observe these variations in the case of each pair of differentiating characters, and to deduce the law according to which they appear in the successive generations. The experiment resolves itself therefore into just as many separate experiments as there are constantly differentiating characters presented in the experimental plants.

The various forms of Peas selected for crossing showed differences in the length and colour of the stem; in the size and form of the leaves; in the position, colour, and size of the flowers; in the length of the flower stalk; in the colour, form, and size of the pods; in the form and size of the seeds; and in the colour of the seed-coats and the albumen [cotyledons]. Some of the characters noted do not permit of a sharp and certain separation, since the difference is of a “more or less” nature, which is often difficult to define. Such characters could not be utilised for the separate experiments; these could only be confined to characters which stand out clearly and definitely in the plants. Lastly, the result must show whether they, in their entirety, observe a regular behaviour in their hybrid unions, and whether from these facts any conclusion can be come to regarding those characters which possess a subordinate significance in the type.

The characters which were selected for experiment relate:

1. To the difference in the form of the ripe seeds. These are either round or roundish, the wrinkling, when such occurs on the surface, being always only shallow; or they are irregularly angular and deeply wrinkled (P. quadratum).

2. To the difference in the colour of the seed albumen (endosperm)[25]. The albumen of the ripe seeds is either pale yellow, bright yellow and orange coloured, or it possesses a more or less intense green tint. This difference of colour is easily seen in the seeds as their coats are transparent.

3. To the difference in the colour of the seed-coat. This is either white, with which character white flowers are constantly correlated; or it is grey, grey-brown, leather-brown, with or without violet spotting, in which case the colour of the standards is violet, that of the wings purple, and the stem in the axils of the leaves is of a reddish tint. The grey seed-coats become dark brown in boiling water.

4. To the difference in the form of the ripe pods. These are either simply inflated, never contracted in places; or they are deeply constricted between the seeds and more or less wrinkled (P. saccharatum).

5. To the difference in the colour of the unripe pods. They are either light to dark green, or vividly yellow, in which colouring the stalks, leaf-veins, and calyx participate[26].

6. To the difference in the position of the flowers. They are either axial, that is, distributed along the main stem; or they are terminal, that is, bunched at the top of the stem and arranged almost in a false umbel; in this case the upper part of the stem is more or less widened in section (P. umbellatum)[27].

7. To the difference in the length of the stem. The length of the stem[28] is very various in some forms; it is, however, a constant character for each, in so far that healthy plants, grown in the same soil, are only subject to unimportant variations in this character.

In experiments with this character, in order to be able to discriminate with certainty, the long axis of 6–7 ft. was always crossed with the short one of 3/4 ft. to 1 1/2 ft.

Each two of the differentiating characters enumerated above were united by cross-fertilisation. There were made for the

1st

trial

fertilisations

plants.

2nd

3rd

4th

5th

6th

7th

From a larger number of plants of the same variety only the most vigorous were chosen for fertilisation. Weakly plants always afford uncertain results, because even in the first generation of hybrids, and still more so in the subsequent ones, many of the offspring either entirely fail to flower or only form a few and inferior seeds.

Furthermore, in all the experiments reciprocal crossings were effected in such a way that each of the two varieties which in one set of fertilisations served as seed-bearers in the other set were used as pollen plants.

The plants were grown in garden beds, a few also in pots, and were maintained in their naturally upright position by means of sticks, branches of trees, and strings stretched between. For each experiment a number of pot plants were placed during the blooming period in a greenhouse, to serve as control plants for the main experiment in the open as regards possible disturbance by insects. Among the insects[29] which visit Peas the beetle Bruchus pisi might be detrimental to the experiments should it appear in numbers. The female of this species is known to lay the eggs in the flower, and in so doing opens the keel; upon the tarsi of one specimen, which was caught in a flower, some pollen grains could clearly be seen under a lens. Mention must also be made of a circumstance which possibly might lead to the introduction of foreign pollen. It occurs, for instance, in some rare cases that certain parts of an otherwise quite normally developed flower wither, resulting in a partial exposure of the fertilising organs. A defective development of the keel has also been observed, owing to which the stigma and anthers remained partially uncovered[30]. It also sometimes happens that the pollen does not reach full perfection. In this event there occurs a gradual lengthening of the pistil during the blooming period, until the stigmatic tip protrudes at the point of the keel. This remarkable appearance has also been observed in hybrids of Phaseolus and Lathyrus.

The risk of false impregnation by foreign pollen is, however, a very slight one with Pisum, and is quite incapable of disturbing the general result. Among more than 10,000 plants which were carefully examined there were only a very few cases where an indubitable false impregnation had occurred. Since in the greenhouse such a case was never remarked, it may well be supposed that Bruchus pisi, and possibly also the described abnormalities in the floral structure, were to blame.

The Forms of the Hybrids.[31]

Experiments which in previous years were made with ornamental plants have already afforded evidence that the hybrids, as a rule, are not exactly intermediate between the parental species. With some of the more striking characters, those, for instance, which relate to the form and size of the leaves, the pubescence of the several parts, &c., the intermediate, indeed, was nearly always to be seen; in other cases, however, one of the two parental characters was so preponderant that it was difficult, or quite impossible, to detect the other in the hybrid.

This is precisely the case with the Pea hybrids. In the case of each of the seven crosses the hybrid-character resembles[32] that of one of the parental forms so closely that the other either escapes observation completely or cannot be detected with certainty. This circumstance is of great importance in the determination and classification of the forms under which the offspring of the hybrids appear. Henceforth in this paper those characters which are transmitted entire, or almost unchanged in the hybridisation, and therefore in themselves constitute the characters of the hybrid, are termed the dominant, and those which become latent in the process recessive. The expression “recessive” has been chosen because the characters thereby designated withdraw or entirely disappear in the hybrids, but nevertheless reappear unchanged in their progeny, as will be demonstrated later on.

It was furthermore shown by the whole of the experiments that it is perfectly immaterial whether the dominant character belong to the seed-bearer or to the pollen parent; the form of the hybrid remains identical in both cases. This interesting fact was also emphasised by Gärtner, with the remark that even the most practised expert is not in a position to determine in a hybrid which of the two parental species was the seed or the pollen plant[33].

Of the differentiating characters which were used in the experiments the following are dominant:

1. The round or roundish form of the seed with or without shallow depressions.

2. The yellow colouring of the seed albumen [cotyledons].

3. The grey, grey-brown, or leather-brown colour of the seed-coat, in connection with violet-red blossoms and reddish spots in the leaf axils.

4. The simply inflated form of the pod.

5. The green colouring of the unripe pod in connection with the same colour in the stems, the leaf-veins and the calyx.

6. The distribution of the flowers along the stem.

7. The greater length of stem.

With regard to this last character it must be stated that the longer of the two parental stems is usually exceeded by the hybrid, which is possibly only attributable to the greater luxuriance which appears in all parts of plants when stems of very different length are crossed. Thus, for instance, in repeated experiments, stems of 1 ft. and 6 ft. in length yielded without exception hybrids which varied in length between 6 ft. and 7 1/2 ft.

The hybrid seeds in the experiments with seed-coat are often more spotted, and the spots sometimes coalesce into small bluish-violet patches. The spotting also frequently appears even when it is absent as a parental character.

The hybrid forms of the seed-shape and of the albumen are developed immediately after the artificial fertilisation by the mere influence of the foreign pollen. They can, therefore, be observed even in the first year of experiment, whilst all the other characters naturally only appear in the following year in such plants as have been raised from the crossed seed.