So recurring to the simile of the man as made by the mixing of tinctures, the process of redistribution of his characters among the germ-cells may be represented as a sorting back of the tinctures again into a double row of bottles, a pair corresponding to each ingredient; and each of the germ-cells as then made of a drop from one or other bottle of each pair: and in our model we may represent the phenomenon of segregation in a crude way by supposing that the bottles having no tincture in them, instead of being empty contained an inoperative fluid, say water, with which the tincture would not mix. When the new germ-cells are formed, the two fluids instead of diluting each other simply separate again. It is this fact which entitles us to speak of the purity of germ-cells. They are pure in the possession of an ingredient, or in not possessing it; and the ingredients, or factors, as we generally call them, are units because they are so treated in the process of formation of the new gametes and because they come out of the process of segregation in the same condition as they went in at fertilisation.
As a consequence of these facts it follows that however complex may be the origin of two given parents the composition of the offspring they can produce is limited. There is only a limited number of types to be made by the possible recombinations of the parental ingredients, and the relative numbers in which each type will be represented are often predicable by very simple arithmetical rules.
For example, if neither parent possesses a certain factor at all, then none of the offspring will have it. If either parent has two doses of the factor then all the children will have it; and if either parent has one dose of the factor and the other has none, then on an average half the family will have it, and half be without it.
To know whether the parent possesses the factor or not may be difficult for reasons which will presently appear, but often it is quite easy and can be told at once, for there are many factors which cannot be present in the individual without manifesting their presence. I may illustrate the descent of such a factor by the case of a family possessing a peculiar form of night-blindness. The affected individuals marrying with those unaffected have a mixture of affected and unaffected children, but their unaffected children not having the responsible ingredient cannot pass it on[1].
In such an observation two things are strikingly exemplified, (1) the fact of the permanence of the unit, and (2) the fact that a mixture of types in the family means that one or other parent is cross-bred in some respect, and is giving off gametes of more than one type.
The problem of heredity is thus a problem primarily analytical. We have to detect and enumerate the factors out of which the bodies of animals and plants are built up, and the laws of their distribution among the germ-cells. All the processes of which I have spoken are accomplished by means of cell-divisions, and in the one cell-union which occurs in fertilisation. If we could watch the factors segregating from each other in cell-division, or even if by microscopic examination we could recognize this multitudinous diversity of composition that must certainly exist among the germ-cells of all ordinary individuals, the work of genetics would be much simpler than it is.
But so far no such direct method of observation has been discovered. In default we are obliged to examine the constitution of the germ-cells by experimental breeding, so contrived that each mating shall test the composition of an individual in one or more chosen respects, and, so to speak, sample its germ-cells by counting the number of each kind of offspring which it can produce. But cumbersome as this method must necessarily be, it enables us to put questions to Nature which never have been put before. She, it has been said, is an unwilling witness. Our questions must be shaped in such a way that the only possible answer is a direct "Yes" or a direct "No." By putting such questions we have received some astonishing answers which go far below the surface. Amazing though they be, they are nevertheless true; for though our witness may prevaricate, she cannot lie. Piecing these answers together, getting one hint from this experiment, and another from that, we begin little by little to reconstruct what is going on in that hidden world of gametes. As we proceed, like our brethren in other sciences, we sometimes receive answers which seem inconsistent or even contradictory. But by degrees a sufficient body of evidence can be attained to show what is the rule and what the exception. My purpose today must be to speak rather of the regular than of the irregular.
One clear exception I may mention. Castle finds that in a cross between the long-eared lop-rabbit and a short-eared breed, ears of intermediate length are produced: and that these intermediates breed approximately true.
Exceptions in general must be discussed elsewhere. Nevertheless if I may throw out a word of counsel to beginners, it is: Treasure your exceptions! When there are none, the work gets so dull that no one cares to carry it further. Keep them always uncovered and in sight. Exceptions are like the rough brickwork of a growing building which tells that there is more to come and shows where the next construction is to be.