Mendel's principles of heredity: A defence

by William Bateson; Gregor Mendel

The First Generation [Bred] from the Hybrids.

In this generation there reappear, together with the dominant characters, also the recessive ones with their full peculiarities, and this occurs in the definitely expressed average proportion of three to one, so that among each four plants of this generation three display the dominant character and one the recessive. This relates without exception to all the characters which were embraced in the experiments. The angular wrinkled form of the seed, the green colour of the albumen, the white colour of the seed-coats and the flowers, the constrictions of the pods, the yellow colour of the unripe pod, of the stalk of the calyx, and of the leaf venation, the umbel-like form of the inflorescence, and the dwarfed stem, all reappear in the numerical proportion given without any essential alteration. Transitional forms were not observed in any experiment.

Once the hybrids resulting from reciprocal crosses are fully formed, they present no appreciable difference in their subsequent development, and consequently the results [of the reciprocal crosses] can be reckoned together in each experiment. The relative numbers which were obtained for each pair of differentiating characters are as follows:

Expt. 1. Form of seed.—From 253 hybrids 7,324 seeds were obtained in the second trial year. Among them were 5,474 round or roundish ones and 1,850 angular wrinkled ones. Therefrom the ratio 2·96 to 1 is deduced.

Expt. 2. Colour of albumen.—258 plants yielded 8,023 seeds, 6,022 yellow, and 2,001 green; their ratio, therefore, is as 3·01 to 1.

In these two experiments each pod yielded usually both kinds of seed. In well-developed pods which contained on the average six to nine seeds, it often occurred that all the seeds were round (Expt. 1) or all yellow (Expt. 2); on the other hand there were never observed more than five angular or five green ones in one pod. It appears to make no difference whether the pods are developed early or later in the hybrid or whether they spring from the main axis or from a lateral one. In some few plants only a few seeds developed in the first formed pods, and these possessed exclusively one of the two characters, but in the subsequently developed pods the normal proportions were maintained nevertheless.

As in separate pods, so did the distribution of the characters vary in separate plants. By way of illustration the first ten individuals from both series of experiments may serve[34].

	Experiment 1.		Experiment 2.
	Form of Seed.		Colour of Albumen.
Plants.	Round.	Angular.	Yellow.	Green.
1	45	12	25	11
2	27	8	32	7
3	24	7	14	5
4	19	10	70	27
5	32	11	24	13
6	26	6	20	6
7	88	24	32	13
8	22	10	44	9
9	28	6	50	14
10	25	7	44	18

As extremes in the distribution of the two seed characters in one plant, there were observed in Expt. 1 an instance of 43 round and only 2 angular, and another of 14 round and 15 angular seeds. In Expt. 2 there was a case of 32 yellow and only 1 green seed, but also one of 20 yellow and 19 green.

These two experiments are important for the determination of the average ratios, because with a smaller number of experimental plants they show that very considerable fluctuations may occur. In counting the seeds, also, especially in Expt. 2, some care is requisite, since in some of the seeds of many plants the green colour of the albumen is less developed, and at first may be easily overlooked. The cause of the partial disappearance of the green colouring has no connection with the hybrid-character of the plants, as it likewise occurs in the parental variety. This peculiarity is also confined to the individual and is not inherited by the offspring. In luxuriant plants this appearance was frequently noted. Seeds which are damaged by insects during their development often vary in colour and form, but, with a little practice in sorting, errors are easily avoided. It is almost superfluous to mention that the pods must remain on the plants until they are thoroughly ripened and have become dried, since it is only then that the shape and colour of the seed are fully developed.

Expt. 3. Colour of the seed-coats.—Among 929 plants 705 bore violet-red flowers and grey-brown seed-coats; 224 had white flowers and white seed-coats, giving the proportion 3·15 to 1.

Expt. 4. Form of pods.—Of 1,181 plants 882 had them simply inflated, and in 299 they were constricted. Resulting ratio, 2·95 to 1.

Expt. 5. Colour of the unripe pods.—The number of trial plants was 580, of which 428 had green pods and 152 yellow ones. Consequently these stand in the ratio 2·82 to 1.

Expt. 6. Position of flowers.—Among 858 cases 651 blossoms were axial and 207 terminal. Ratio, 3·14 to 1.

Expt. 7. Length of stem.—Out of 1,064 plants, in 787 cases the stem was long, and in 277 short. Hence a mutual ratio of 2·84 to 1. In this experiment the dwarfed plants were carefully lifted and transferred to a special bed. This precaution was necessary, as otherwise they would have perished through being overgrown by their tall relatives. Even in their quite young state they can be easily picked out by their compact growth and thick dark-green foliage.

If now the results of the whole of the experiments be brought together, there is found, as between the number of forms with the dominant and recessive characters, an average ratio of 2·98 to 1, or 3 to 1.

The dominant character can have here a double signification—viz. that of a parental-character, or a hybrid-character[35]. In which of the two significations it appears in each separate case can only be determined by the following generation. As a parental character it must pass over unchanged to the whole of the offspring; as a hybrid-character, on the other hand, it must observe the same behaviour as in the first generation.