Starting from this standpoint the relative functions of the two sexes in heredity are apparent. The original function of reproduction, that of cell division, is the part of the female. The male in the lower instances simply supplies the female with nutriment. Thus in certain plants there is nothing but a subtle osmosis between the sexes. This is also the case with some of the lower infusoria. With a rise in evolution protoplasm becomes differentiated. At the outset of the subject of heredity it is evident, therefore, that the female furnishes the type which is best capable of development when properly nourished by a highly developed male. To deficiencies in both particulars are due defects and variations in the offspring. As the product of fructification is longest under the nutritive control of the female, her influence is most emphatic in either redeeming defects or producing them. Heredity, according to Ribot, Spitzka, Féré, and others, is divisible into direct heredity, indirect heredity, and, more dubiously, telegony. Direct heredity consists in the transmission of paternal and maternal qualities to the children. This form of heredity has two aspects: (1) The child takes after father and mother equally as regards both physical and moral characters, a case strictly speaking of very rare occurrence; or (2) the child, while taking after both parents, more especially resembles one of them. Here again distinction must be made between two cases. The first of these is when the heredity takes place in the same sex from father to son, from mother to daughter. The other which occurs more frequently, is where heredity occurs between different sexes—from father to daughter or from mother to son. Reversional heredity or atavism consists in the reproduction in the descendants of the moral or physical qualities of their ancestors. It occurs frequently between grandfather and grandson, as well as between grandmother and granddaughter. Collateral or indirect heredity, which is of rarer occurrence than the foregoing, and is simply a form of atavism, subsists, as indicated by the name, between individuals and their ancestors in the indirect line—uncle, or grand-uncle and nephew, aunt and niece. Finally (3) there is telegony, or the heredity of influence, very rare from the physiological point of view, which consists in reproduction in the children by a second marriage of some peculiarity belonging to a former spouse.
In dealing with heredity the position of Weismann and others, that acquired characters cannot be inherited, needs a short examination. In his later work Weismann has practically abandoned the essential basis of his position by admitting that maternal nutrition may play a part in determining variation. He[68] now asserts that the origin of a variation is equally independent of selection and amphimixis, and is due to the constant occurrence of slight inequalities of nutrition in the germ plasm. As acquired characters affecting the constitution of the parents are certain to affect the nutrition of the germ plasm, it is therefore obvious, according to Weismann’s admission, that acquired characters or their consequences will be inherited. This is an emphatic though concealed abandonment of the central position of Weismann.
One of the stock arguments of the Weismann school is drawn from results of the Jewish rite of circumcision. While the operation is not calculated to make a profound impression on the constitution, and furthermore, as being performed on the male, less likely to affect the race, still the alleged non-inheritance of its results is much over-estimated. William Wolf,[69] of Baltimore, Maryland, who has circumcised six hundred Jewish children, finds on careful examination, that 2 per cent. were born partially circumcised and 6 per cent. were born with a short prepuce. P. C. Remondino,[70] of Los Angeles, California, has seen a large number of cases of absence of the prepuce which proved to be hereditary. After a confinement his attention was once called to the child by the nurse, who thought it was deformed. The nurse was astonished at the size and appearance of the glans penis. On examination the prepuce was found to be completely absent. On inquiry, the father and another son, born more than twenty years previously (comprising every male member of the family), were found to have been born with the glands fully exposed. He has seen a French family similarly affected.
Similar, but much stronger, results have been obtained by me through the courtesy of the Reverend Drs. S. Bauer, M. A. Cohen, and B. Gordon, all of Chicago. Dr. Bauer, who has been seventeen years in the practice of the religious rite of circumcision, has circumcised 3,400 boys and has found preputial absence in about 3½ per cent. of the cases. Dr. Cohen, who has been two decades in the practice, has performed 10,000 circumcisions. He has found the prepuce wanting in 500 cases; partially developed in 300 cases; slightly developed in 2,000 cases. Dr. Gordon has performed 4,400 circumcisions in twenty-five years. He has found the prepuce absent in 15 cases; partly wanting in 200, and slightly developed in 2,200 cases. These, it should be remembered, are only cases where preputial change forced itself on the observer, who was not pursuing investigations on this point.
The volume of Hebrew casuistic religious literature collected in the Medrash, evidences as I have elsewhere[71] shown the frequency of congenital preputial defect.
That acquired characters can be transmitted has been definitely shown by the experiments of George Roe Lockwood,[72] of New York, anent hereditary transmission of mutilations. White mice were selected, as they begin to breed when thirty days old, and breed every thirty days. He bred in-and-in for thirty-six generations, destroying the weakly, and thus obtained finer animals than the first pair. He selected a pair, caged them by themselves, and clipped the tails of the young. When they were old enough to breed, he selected a pair and clipped the tails of their progeny. In the seventh generation he obtained some tailless mice, and finally a tailless breed. The experiments have one possible element of error; white mice, like all albinoes, are a degenerate type. At the same time these experiments show that accidental mutilations favoured by circumstances are inherited.
Eimer[73] reports the case of a pair of long-tailed pointers which had once produced a litter of long-tailed pups. In order to obtain short-tailed pups the owner had the tails of both shortened. The bitch from that time produced repeatedly short-tailed pups only. As the most careful attention was paid to the parents, no error can be suspected in this case, which, moreover, excites no surprise among dog-breeders.
Brown-Séquard[74] has shown that a peculiar alteration of the shape of the ear or a partial closing of the eyelids is inherited by the offspring of animals in which these changes were caused by dividing the sympathetic. Exophthalmia (eye protrusion) was inherited by guinea-pigs in whose parents this protrusion of the eyes had occurred after an injury to the spinal cord, and so were bruises and dry gangrene, as well as other trophic disturbances in the ear, due in the parents to an injury to the restiform body of the brain. Loss of certain phalanges or of whole toes of the hind feet which had occurred in the parents in consequence of division of the sciatic nerve was inherited. Diseased conditions of the sciatic nerve occurred in the offspring of guinea-pigs in which this nerve was divided. Forty guinea-pigs in which one or both eyes showed more or less morbid change were descended from three individuals in which one eye had become diseased in consequence of transverse section of the restiform body. Twenty guinea-pigs exhibited muscular atrophy on the upper and lower sides of the thigh, when in the parents such atrophy had been caused by section of the sciatic nerve.
The experiments of Brown-Séquard, Westphal, Dupuy[75] and Obersteiner, which show that artificially induced epilepsy is inherited, still further bear out the conclusions resultant on the inheritance of these mutilations. Indeed, Weismann has been forced to that reductio ad absurdum in science, narrowly limited definitions, in order to maintain his position. “But although I hold it improbable,” he remarks, “that individual variability can depend on a direct action of external influences upon the germ cells and their contained germ plasm, because, as follows from sundry facts, the molecular structure of the germ plasm must be very difficult to change, yet it is by no means to be implied that this structure may not possibly be altered by influences of the same kind continuing for a very long time. This much may be maintained: that influences which are mostly of variable nature, tending now in one direction, now in another, can hardly produce a change in the structure of the germ plasm, and this is the reason why the cause of inheritable individual differences must be sought elsewhere than in these varying influences.” “No one has doubted,” he says, in reply to objections made by Virchow, “that there are a number of congenital deformities, birth-marks, and other individual peculiarities which are inherited. But these are acquired characters in the above sense. True, they must once have appeared for the first time, but we cannot say exactly from what causes; we only know that at least a great proportion of them proceed from the germ itself, and must therefore be due to alterations of the germinal substance. If Virchow could show that any single one of these hereditary deformities had its origin in the action of some external cause upon the already formed body (soma) of the individual and not upon the germ cell, then the inheritance of acquired characters would be proved. But this no one has yet succeeded in proving, often as it has been maintained.”
The crucial test which Weismann demands is furnished by Dupuy,[76] who made one thousand experiments on guinea-pigs to the fifth generation, critically rejecting all results which did not correspond to the most rigid tests of direct heredity, excluding all instances of indirect heredity, however demonstrable. He found that certain lesions of the spinal cord, or the brain or the sciatic nerve, give rise in guinea-pigs to epilepsy.