The hyacinth delights in a rich light sandy soil. The Dutch incorporate freely with their naturally light soil a compost consisting of one-third coarse sea or river sand, one-third rotten cow dung without litter and one-third leaf-mould. The soil thus renovated retains its qualities for six or seven years, but hyacinths are not planted upon the same place for two years successively, intermediary crops of narcissus, crocus or tulips being taken. A good compost for hyacinths is sandy loam, decayed leaf-mould, rotten cow dung and sharp sand in equal parts, the whole being collected and laid up in a heap and turned over occasionally. Well-drained beds made up of this soil, and refreshed with a portion of new compost annually, would grow the hyacinth to perfection. The best time to plant the bulbs is towards the end of September and during October; they should be arranged in rows, 6 to 8 in. asunder, there being four rows in each bed. The bulbs should be sunk about 4 to 6 in. deep, with a small quantity of clean sand placed below and around each of them. The beds should be covered with decayed tan-bark, coco-nut fibre or half-rotten dung litter. As the flower-stems appear, they are tied to rigid but slender stakes to preserve them from accident. If the bulbs are at all prized, the stems should be broken off as soon as the flowering is over, so as not to exhaust the bulbs; the leaves, however, must be allowed to grow on till matured, but as soon as they assume a yellow colour, the bulbs are taken up, the leaves cut off near their base, and the bulbs laid out in a dry, airy, shady place to ripen, after which they are cleaned of loose earth and skin, ready for storing. It is the practice in Holland, about a month after the bloom, or when the tips of the leaves assume a withered appearance, to take up the bulbs, and to lay them sideways on the ground, covering them with an inch or two of earth. About three weeks later they are again taken up and cleaned. In the store-room they should be kept dry, well-aired and apart from each other.Few plants are better adapted than the hyacinth for pot culture as greenhouse decorative plants; and by the aid of forcing they may be had in bloom as early as Christmas. They flower fairly well in 5-in. pots, the stronger bulbs in 6-in. pots. To bloom at Christmas, they should be potted early in September, in a compost resembling that already recommended for the open-air beds; and, to keep up a succession of bloom, others should be potted at intervals of a few weeks till the middle or end of November. The tops of the bulbs should be about level with the soil, and if a little sand is put immediately around them so much the better. The pots should be set in an open place on a dry hard bed of ashes, and be covered over to a depth of 6 or 8 in. with the same material or with fibre or soil; and when the roots are well developed, which will take from six to eight weeks, they may be removed to a frame, and gradually exposed to light, and then placed in a forcing pit in a heat of from 60 to 70°. When the flowers are fairly open, they may be removed to the greenhouse or conservatory.The hyacinth may be very successfully grown in glasses for ornament in dwelling-houses. The glasses are filled to the neck with rain or even tap water, a few lumps of charcoal being dropped into them. The bulbs are placed in the hollow provided for them, so that their base just touches the water. This may be done in September or October. They are then set in a dark cupboard for a few weeks till roots are freely produced, and then gradually exposed to light. The early-flowering single white Roman hyacinth, a small-growing pure white variety, remarkable for its fragrance, is well adapted for forcing, as it can be had in bloom if required by November. For windows it grows well in the small glasses commonly used for crocuses; and for decorative purposes should be planted about five bulbs in a 5-in. pot, or in pans holding a dozen each. If grown for cut flowers it can be planted thickly in boxes of any convenient size. It is highly esteemed during the winter months by florists.The Spanish hyacinth (H. amethystinus) andH. azureusare charming little bulbs for growing in masses in the rock garden or front of the flower border. The older botanists included in the genusHyacinthusspecies ofMuscari,Scillaand other genera of bulbous Liliaceae, and the name of hyacinth is still popularly applied to several other bulbous plants. ThusMuscari botryoidesis the grape hyacinth, 6 in., blue or white, the handsomest;M. moschatum, the musk hyacinth, 10 in., has peculiar livid greenish-yellow flowers and a strong musky odour;M. comosumvar.monstrosum, the feather hyacinth, bears sterile flowers broken up into a featherlike mass;M. racemosum, the starch hyacinth, is a native with deep blue plum-scented flowers. The Cape hyacinth isGaltonia candicans, a magnificent border plant, 3-4 ft. high, with large drooping white bell-shaped flowers; the star hyacinth,Scilla amoena; the Peruvian hyacinth or Cuban lily,S. peruviana, a native of the Mediterranean region, to which Linnaeus gave the species nameperuvianaon a mistaken assumption of its origin; the wild hyacinth or blue-bell, known variously asEndymion nonscriptum,Hyacinthus nonscriptusorScilla nutans; the wild hyacinth of western NorthAmerica,Camassia esculenta. They all flourish in good garden soil of a gritty nature.
The hyacinth delights in a rich light sandy soil. The Dutch incorporate freely with their naturally light soil a compost consisting of one-third coarse sea or river sand, one-third rotten cow dung without litter and one-third leaf-mould. The soil thus renovated retains its qualities for six or seven years, but hyacinths are not planted upon the same place for two years successively, intermediary crops of narcissus, crocus or tulips being taken. A good compost for hyacinths is sandy loam, decayed leaf-mould, rotten cow dung and sharp sand in equal parts, the whole being collected and laid up in a heap and turned over occasionally. Well-drained beds made up of this soil, and refreshed with a portion of new compost annually, would grow the hyacinth to perfection. The best time to plant the bulbs is towards the end of September and during October; they should be arranged in rows, 6 to 8 in. asunder, there being four rows in each bed. The bulbs should be sunk about 4 to 6 in. deep, with a small quantity of clean sand placed below and around each of them. The beds should be covered with decayed tan-bark, coco-nut fibre or half-rotten dung litter. As the flower-stems appear, they are tied to rigid but slender stakes to preserve them from accident. If the bulbs are at all prized, the stems should be broken off as soon as the flowering is over, so as not to exhaust the bulbs; the leaves, however, must be allowed to grow on till matured, but as soon as they assume a yellow colour, the bulbs are taken up, the leaves cut off near their base, and the bulbs laid out in a dry, airy, shady place to ripen, after which they are cleaned of loose earth and skin, ready for storing. It is the practice in Holland, about a month after the bloom, or when the tips of the leaves assume a withered appearance, to take up the bulbs, and to lay them sideways on the ground, covering them with an inch or two of earth. About three weeks later they are again taken up and cleaned. In the store-room they should be kept dry, well-aired and apart from each other.
Few plants are better adapted than the hyacinth for pot culture as greenhouse decorative plants; and by the aid of forcing they may be had in bloom as early as Christmas. They flower fairly well in 5-in. pots, the stronger bulbs in 6-in. pots. To bloom at Christmas, they should be potted early in September, in a compost resembling that already recommended for the open-air beds; and, to keep up a succession of bloom, others should be potted at intervals of a few weeks till the middle or end of November. The tops of the bulbs should be about level with the soil, and if a little sand is put immediately around them so much the better. The pots should be set in an open place on a dry hard bed of ashes, and be covered over to a depth of 6 or 8 in. with the same material or with fibre or soil; and when the roots are well developed, which will take from six to eight weeks, they may be removed to a frame, and gradually exposed to light, and then placed in a forcing pit in a heat of from 60 to 70°. When the flowers are fairly open, they may be removed to the greenhouse or conservatory.
The hyacinth may be very successfully grown in glasses for ornament in dwelling-houses. The glasses are filled to the neck with rain or even tap water, a few lumps of charcoal being dropped into them. The bulbs are placed in the hollow provided for them, so that their base just touches the water. This may be done in September or October. They are then set in a dark cupboard for a few weeks till roots are freely produced, and then gradually exposed to light. The early-flowering single white Roman hyacinth, a small-growing pure white variety, remarkable for its fragrance, is well adapted for forcing, as it can be had in bloom if required by November. For windows it grows well in the small glasses commonly used for crocuses; and for decorative purposes should be planted about five bulbs in a 5-in. pot, or in pans holding a dozen each. If grown for cut flowers it can be planted thickly in boxes of any convenient size. It is highly esteemed during the winter months by florists.
The Spanish hyacinth (H. amethystinus) andH. azureusare charming little bulbs for growing in masses in the rock garden or front of the flower border. The older botanists included in the genusHyacinthusspecies ofMuscari,Scillaand other genera of bulbous Liliaceae, and the name of hyacinth is still popularly applied to several other bulbous plants. ThusMuscari botryoidesis the grape hyacinth, 6 in., blue or white, the handsomest;M. moschatum, the musk hyacinth, 10 in., has peculiar livid greenish-yellow flowers and a strong musky odour;M. comosumvar.monstrosum, the feather hyacinth, bears sterile flowers broken up into a featherlike mass;M. racemosum, the starch hyacinth, is a native with deep blue plum-scented flowers. The Cape hyacinth isGaltonia candicans, a magnificent border plant, 3-4 ft. high, with large drooping white bell-shaped flowers; the star hyacinth,Scilla amoena; the Peruvian hyacinth or Cuban lily,S. peruviana, a native of the Mediterranean region, to which Linnaeus gave the species nameperuvianaon a mistaken assumption of its origin; the wild hyacinth or blue-bell, known variously asEndymion nonscriptum,Hyacinthus nonscriptusorScilla nutans; the wild hyacinth of western NorthAmerica,Camassia esculenta. They all flourish in good garden soil of a gritty nature.
HYACINTH,orJacinth, in mineralogy, a variety of zircon (q.v.) of yellowish red colour, used as a gem-stone. Thehyacinthusof ancient writers must have been our sapphire, or blue corundum, while the hyacinth of modern mineralogists may have been the stone known aslyncurium(λυγκούριον). The Hebrew wordleshem, translated ligure in the Authorized Version (Ex. xxviii. 19), from theλιγύριονof the Septuagint, appears in the Revised Version as jacinth, but with a marginal alternative of amber. Both jacinth and amber may be reddish yellow, but their identification is doubtful. As our jacinth (zircon) is not known in ancient Egyptian work, Professor Flinders Petrie has suggested that theleshemmay have been a yellow quartz, or perhaps agate. Some old English writers describe the jacinth as yellow, whilst others refer to it as a blue stone, and thehyacinthusof some authorities seems undoubtedly to have been our sapphire. In Rev. xx. 20 the Revised Version retains the word jacinth, but gives sapphire as an alternative.
Most of the gems known in trade as hyacinth are only garnets—generally the deep orange-brown hessonite or cinnamon-stone—and many of the antique engraved stones reputed to be hyacinth are probably garnets. The difference may be detected optically, since the garnet is singly and the hyacinth doubly refracting; moreover the specific gravity affords a simple means of diagnosis, that of garnet being only about 3.7, whilst hyacinth may have a density as high as 4.7. Again, it was shown many years ago by Sir A. H. Church that most hyacinths, when examined by the spectroscope, show a series of dark absorption bands, due perhaps to the presence of some rare element such as uranium or erbium.
Hyacinth is not a common mineral. It occurs, with other zircons, in the gem-gravels of Ceylon, and very fine stones have been found as pebbles at Mudgee in New South Wales. Crystals of zircon, with all the typical characters of hyacinth, occur at Expailly, Le Puy-en-Velay, in Central France, but they are not large enough for cutting. The stones which have been called Compostella hyacinths are simply ferruginous quartz from Santiago de Compostella in Spain.
(F. W. R.*)
HYACINTHUS,1in Greek mythology, the youngest son of the Spartan king Amyclas, who reigned at Amyclae (so Pausanias iii. 1. 3, iii. 19. 5; and Apollodorus i. 3. 3, iii. 10. 3). Other stories make him son of Oebalus, of Eurotas, or of Pierus and the nymph Clio (see Hyginus,Fabulae, 271; Lucian,De saltatione, 45, andDial. deor.14). According to the general story, which is probably late and composite, his great beauty attracted the love of Apollo, who killed him accidentally when teaching him to throw thediscus(quoit); others say that Zephyrus (or Boreas) out of jealousy deflected the quoit so that it hit Hyacinthus on the head and killed him. According to the representation on the tomb at Amyclae (Pausanias,loc. cit.) Hyacinthus was translated into heaven with his virgin sister Polyboea. Out of his blood there grew the flower known as the hyacinth, the petals of which were marked with the mournful exclamation AI, AI, “alas” (cf. “that sanguine flower inscribed with woe”). This Greek hyacinth cannot have been the flower which now bears the name: it has been identified with a species of iris and with the larkspur (Delphinium Aiacis), which appear to have the markings described. The Greek hyacinth was also said to have sprung from the blood of Ajax. Evidently the Greek authorities confused both the flowers and the traditions.
The death of Hyacinthus was celebrated at Amyclae by the second most important of Spartan festivals, the Hyacinthia, which took place in the Spartan month Hecatombeus. What month this was is not certain. Arguing from Xenophon (Hell.iv. 5) we get May; assuming that the Spartan Hecatombeus is the Attic Hecatombaion, we get July; or again it may be the Attic Scirophorion, June. At all events the Hyacinthia was an early summer festival. It lasted three days, and the rites gradually passed from mourning for Hyacinthus to rejoicingsin the majesty of Apollo, the god of light and warmth, and giver of the ripe fruits of the earth (see a passage from Polycrates,Laconica, quoted by Athenaeus 139 d; criticized by L. R. Farnell,Cults of the Greek States, iv. 266 foll.). This festival is clearly connected with vegetation, and marks the passage from the youthful verdure of spring to the dry heat of summer and the ripening of the corn.
The precise relation which Apollo bears to Hyacinthus is obscure. The fact that at Tarentum a Hyacinthus tomb is ascribed by Polybius to Apollo Hyacinthus (not Hyacinthius) has led some to think that the personalities are one, and that the hero is merely an emanation from the god; confirmation is sought in the Apolline appellationτετράχειρ, alleged by Hesychius to have been used in Laconia, and assumed to describe a composite figure of Apollo-Hyacinthus. Against this theory is the essential difference between the two figures. Hyacinthus is a chthonian vegetation god whose worshippers are afflicted and sorrowful; Apollo, though interested in vegetation, is never regarded as inhabiting the lower world, his death is not celebrated in any ritual, his worship is joyous and triumphant, and finally the Amyclean Apollo is specifically the god of war and song. Moreover, Pausanias describes the monument at Amyclae as consisting of a rude figure of Apollo standing on an altar-shaped base which formed the tomb of Hyacinthus. Into the latter offerings were put for the hero before gifts were made to the god.
On the whole it is probable that Hyacinthus belongs originally to the pre-Dorian period, and that his story was appropriated and woven into their own Apollo myth by the conquering Dorians. Possibly he may be the apotheosis of a pre-Dorian king of Amyclae. J. G. Frazer further suggests that he may have been regarded as spending the winter months in the underworld and returning to earth in the spring when the “hyacinth” blooms. In this case his festival represents perhaps both the Dorian conquest of Amyclae and the death of spring before the ardent heat of the summer sun, typified as usual by thediscus(quoit) with which Apollo is said to have slain him. With the growth of the hyacinth from his blood should be compared the oriental stories of violets springing from the blood of Attis, and roses and anemones from that of Adonis. As a youthful vegetation god, Hyacinthus may be compared with Linus and Scephrus, both of whom are connected with Apollo Agyieus.
See L. R. Farnell,Cults of the Greek States, vol. iv. (1907), pp. 125 foll., 264 foll.; J. G. Frazer,Adonis, Attis, Osiris(1906), bk. ii. ch. 7; S. Wide,Lakonische Kulte, p. 290; E. Rhode,Psyche, 3rd ed. i. 137 foll.; Roscher,Lexikon d. griech. u. röm. Myth.,s.v.“Hyakinthos” (Greve); L. Preller,Griechische Mythol.4th ed. i. 248 foll.
See L. R. Farnell,Cults of the Greek States, vol. iv. (1907), pp. 125 foll., 264 foll.; J. G. Frazer,Adonis, Attis, Osiris(1906), bk. ii. ch. 7; S. Wide,Lakonische Kulte, p. 290; E. Rhode,Psyche, 3rd ed. i. 137 foll.; Roscher,Lexikon d. griech. u. röm. Myth.,s.v.“Hyakinthos” (Greve); L. Preller,Griechische Mythol.4th ed. i. 248 foll.
(J. M. M.)
1The word is probably derived from an Indo-European root, meaning “youthful,” found in Latin, Greek, English and Sanskrit. Some have suggested that the first two letters are fromὕειν, to rain, (cf. Hyades).
1The word is probably derived from an Indo-European root, meaning “youthful,” found in Latin, Greek, English and Sanskrit. Some have suggested that the first two letters are fromὕειν, to rain, (cf. Hyades).
HYADES(“the rainy ones”), in Greek mythology, the daughters of Atlas and Aethra; their number varies between two and seven. As a reward for having brought up Zeus at Dodona and taken care of the infant Dionysus Hyes, whom they conveyed to Ino (sister of his mother Semele) at Thebes when his life was threatened by Lycurgus, they were translated to heaven and placed among the stars (Hyginus,Poët. astron.ii. 21). Another form of the story combines them with the Pleiades. According to this they were twelve (or fifteen) sisters, whose brother Hyas was killed by a snake while hunting in Libya (Ovid,Fasti, v. 165; Hyginus,Fab.192). They lamented him so bitterly that Zeus, out of compassion, changed them into stars—five into the Hyades, at the head of the constellation of the Bull, the remainder into the Pleiades. Their name is derived from the fact that the rainy season commenced when they rose at the same time as the sun (May 7-21); the original conception of them is that of the fertilizing principle of moisture. The Romans derived the name fromὗς(pig), and translated it bySuculae(Cicero,De nat. deorum, ii. 43).
HYATT, ALPHEUS(1838-1902), American naturalist, was born at Washington, D.C., on the 5th of April 1838. From 1858 to 1862 he studied at Harvard, where he had Louis Agassiz for his master, and in 1863 he served as a volunteer in the Civil War, attaining the rank of captain. In 1867 he was appointed curator of the Essex Institute at Salem, and in 1870 became professor of zoology and palaeontology at the Massachusetts Institute of Technology (resigned 1888), and custodian of the Boston Society of Natural History (curator in 1881). In 1886 he was appointed assistant for palaeontology in the Cambridge museum of comparative anatomy, and in 1889 was attached to the United States Geological Survey as palaeontologist for the Trias and Jura. He was the chief founder of the American Society of Naturalists, of which he acted as first president in 1883, and he also took a leading part in establishing the marine biological laboratories at Annisquam and Woods Hole, Mass. He died at Cambridge on the 15th of January 1902.
His works includeObservations on Fresh-water Polyzoa(1866);Fossil Cephalopods of the Museum of Comparative Zoology(1872);Revision of North American Porifera(1875-1877);Genera of Fossil Cephalopoda(1883);Larval Theory of the Origin of Cellular Tissue(1884);Genesis of the Arietidae(1889); andPhylogeny of an acquired characteristic(1894). He wrote the section on Cephalopoda in Karl von Zittel’sPaläontologie(1900), and his well-known study on the fossil pond snails of Steinheim (“The Genesis of the Tertiary Species of Planorbis at Steinheim”) appeared in theMemoirsof the Boston Natural History Society in 1880. He was one of the founders and editors of theAmerican Naturalist.
His works includeObservations on Fresh-water Polyzoa(1866);Fossil Cephalopods of the Museum of Comparative Zoology(1872);Revision of North American Porifera(1875-1877);Genera of Fossil Cephalopoda(1883);Larval Theory of the Origin of Cellular Tissue(1884);Genesis of the Arietidae(1889); andPhylogeny of an acquired characteristic(1894). He wrote the section on Cephalopoda in Karl von Zittel’sPaläontologie(1900), and his well-known study on the fossil pond snails of Steinheim (“The Genesis of the Tertiary Species of Planorbis at Steinheim”) appeared in theMemoirsof the Boston Natural History Society in 1880. He was one of the founders and editors of theAmerican Naturalist.
HYBLA,the name of several cities In Sicily. The best known historically, though its exact site is uncertain, is Hybla Major, near (or by some supposed to be identical with) Megara Hyblaea (q.v.): another Hybla, known as Hybla Minor or Galeatis, is represented by the modern Paternò; while the site of Hybla Heraea is to be sought near Ragusa.
HYBRIDISM.The Latin wordhybrida,hibridaoribridahas been assumed to be derived from the Greekὕβρις, an insult or outrage, and a hybrid or mongrel has been supposed to be an outrage on nature, an unnatural product. As a general rule animals and plants belonging to distinct species do not produce offspring when crossed with each other, and the term hybrid has been employed for the result of a fertile cross between individuals of different species, the word mongrel for the more common result of the crossing of distinct varieties. A closer scrutiny of the facts, however, makes the term hybridism less isolated and more vague. The words species and genus, and still more subspecies and variety, do not correspond with clearly marked and sharply defined zoological categories, and no exact line can be drawn between the various kinds of crossings from those between individuals apparently identical to those belonging to genera universally recognized as distinct. Hybridism therefore grades into mongrelism, mongrelism into cross-breeding, and cross-breeding into normal pairing, and we can say little more than that the success of the union is the more unlikely or more unnatural the further apart the parents are in natural affinity.
The interest in hybridism was for a long time chiefly of a practical nature, and was due to the fact that hybrids are often found to present characters somewhat different from those of either parent. The leading facts have been known in the case of the horse and ass from time immemorial. The earliest recorded observation of a hybrid plant is by J. G. Gmelin towards the end of the 17th century; the next is that of Thomas Fairchild, who in the second decade of the 18th century, produced the cross which is still grown in gardens under the name of “Fairchild’s Sweet William.” Linnaeus made many experiments in the cross-fertilization of plants and produced several hybrids, but Joseph Gottlieb Kölreuter (1733-1806) laid the first real foundation of our scientific knowledge of the subject. Later on Thomas Andrew Knight, a celebrated English horticulturist, devoted much successful labour to the improvement of fruit trees and vegetables by crossing. In the second quarter of the 19th century C. F. Gärtner made and published the results of a number of experiments that had not been equalled by any earlier worker. Next came Charles Darwin, who first in theOrigin of Species, and later inCross and Self-Fertilization of Plants, subjected the whole question to a critical examination, reviewed the known facts and added many to them.
Darwin’s conclusions were summed up by G. J. Romanes in the 9th edition of thisEncyclopaediaas follows:—1. The laws governing the production of hybrids are identical, or nearly identical, in the animal and vegetable kingdoms.2. The sterility which so generally attends the crossing of two specific forms is to be distinguished as of two kinds, which, althoughoften confounded by naturalists, are in reality quite distinct. For the sterility may obtain between the two parent species when first crossed, or it may first assert itself in their hybrid progeny. In the latter case the hybrids, although possibly produced without any appearance of infertility on the part of their parent species, nevertheless prove more or less infertile among themselves, and also with members of either parent species.3. The degree of both kinds of infertility varies in the case of different species, and in that of their hybrid progeny, from absolute sterility up to complete fertility. Thus, to take the case of plants, “when pollen from a plant of one family is placed on the stigma of a plant of a distinct family, it exerts no more influence than so much inorganic dust. From this absolute zero of fertility, the pollen of different species, applied to the stigma of some one species of the same genus, yields a perfect gradation in the number of seeds produced, up to nearly complete, or even quite complete, fertility; so, in hybrids themselves, there are some which never have produced, and probably never would produce, even with the pollen of the pure parents, a single fertile seed; but in some of these cases a first trace of fertility may be detected, by the pollen of one of the pure parent species causing the flower of the hybrid to wither earlier than it otherwise would have done; and the early withering of the flower is well known to be a sign of incipient fertilization. From this extreme degree of sterility we have self-fertilized hybrids producing a greater and greater number of seeds up to perfect fertility.”4. Although there is, as a rule, a certain parallelism, there is no fixed relation between the degree of sterility manifested by the parent species when crossed and that which is manifested by their hybrid progeny. There are many cases in which two pure species can be crossed with unusual facility, while the resulting hybrids are remarkably sterile; and, contrariwise, there are species which can only be crossed with extreme difficulty, though the hybrids, when produced, are very fertile. Even within the limits of the same genus, these two opposite cases may occur.5. When two species are reciprocally crossed,i.e.male A with female B, and male B with female A, the degree of sterility often differs greatly in the two cases. The sterility of the resulting hybrids may differ likewise.6. The degree of sterility of first crosses and of hybrids runs, to a certain extent, parallel with the systematic affinity of the forms which are united. “For species belonging to distinct genera can rarely, and those belonging to distinct families can never, be crossed. The parallelism, however, is far from complete; for a multitude of closely allied species will not unite, or unite with extreme difficulty, whilst other species, widely different from each other, can be crossed with perfect facility. Nor does the difficulty depend on ordinary constitutional differences; for annual and perennial plants, deciduous and evergreen trees, plants flowering at different seasons, inhabiting different stations, and naturally living under the most opposite climates, can often be crossed with ease. The difficulty or facility apparently depends exclusively on the sexual constitution of the species which are crossed, or on their sexual elective affinity.”
Darwin’s conclusions were summed up by G. J. Romanes in the 9th edition of thisEncyclopaediaas follows:—
1. The laws governing the production of hybrids are identical, or nearly identical, in the animal and vegetable kingdoms.
2. The sterility which so generally attends the crossing of two specific forms is to be distinguished as of two kinds, which, althoughoften confounded by naturalists, are in reality quite distinct. For the sterility may obtain between the two parent species when first crossed, or it may first assert itself in their hybrid progeny. In the latter case the hybrids, although possibly produced without any appearance of infertility on the part of their parent species, nevertheless prove more or less infertile among themselves, and also with members of either parent species.
3. The degree of both kinds of infertility varies in the case of different species, and in that of their hybrid progeny, from absolute sterility up to complete fertility. Thus, to take the case of plants, “when pollen from a plant of one family is placed on the stigma of a plant of a distinct family, it exerts no more influence than so much inorganic dust. From this absolute zero of fertility, the pollen of different species, applied to the stigma of some one species of the same genus, yields a perfect gradation in the number of seeds produced, up to nearly complete, or even quite complete, fertility; so, in hybrids themselves, there are some which never have produced, and probably never would produce, even with the pollen of the pure parents, a single fertile seed; but in some of these cases a first trace of fertility may be detected, by the pollen of one of the pure parent species causing the flower of the hybrid to wither earlier than it otherwise would have done; and the early withering of the flower is well known to be a sign of incipient fertilization. From this extreme degree of sterility we have self-fertilized hybrids producing a greater and greater number of seeds up to perfect fertility.”
4. Although there is, as a rule, a certain parallelism, there is no fixed relation between the degree of sterility manifested by the parent species when crossed and that which is manifested by their hybrid progeny. There are many cases in which two pure species can be crossed with unusual facility, while the resulting hybrids are remarkably sterile; and, contrariwise, there are species which can only be crossed with extreme difficulty, though the hybrids, when produced, are very fertile. Even within the limits of the same genus, these two opposite cases may occur.
5. When two species are reciprocally crossed,i.e.male A with female B, and male B with female A, the degree of sterility often differs greatly in the two cases. The sterility of the resulting hybrids may differ likewise.
6. The degree of sterility of first crosses and of hybrids runs, to a certain extent, parallel with the systematic affinity of the forms which are united. “For species belonging to distinct genera can rarely, and those belonging to distinct families can never, be crossed. The parallelism, however, is far from complete; for a multitude of closely allied species will not unite, or unite with extreme difficulty, whilst other species, widely different from each other, can be crossed with perfect facility. Nor does the difficulty depend on ordinary constitutional differences; for annual and perennial plants, deciduous and evergreen trees, plants flowering at different seasons, inhabiting different stations, and naturally living under the most opposite climates, can often be crossed with ease. The difficulty or facility apparently depends exclusively on the sexual constitution of the species which are crossed, or on their sexual elective affinity.”
There are many new records as to the production of hybrids. Horticulturists have been extremely active and successful in their attempts to produce new flowers or new varieties of vegetables by seminal or graft-hybrids, and any florist’s catalogue or the account of any special plant, such as is to be found in Foster-Melliar’sBook of the Rose, is in great part a history of successful hybridization. Much special experimental work has been done by botanists, notably by de Vries, to the results of whose experiments we shall recur. Experiments show clearly that the obtaining of hybrids is in many cases merely a matter of taking sufficient trouble, and the successful crossing of genera is not infrequent.
Focke, for instance, cites cases where hybrids were obtained betweenBrassicaandRaphanus,GaliumandAsperula,CampanulaandPhyteuma,VerbascumandCelsia. Among animals, new records and new experiments are almost equally numerous. Boveri has crossedEchinus microtuberculatuswithSphaerechinus granularis. Thomas Hunt Morgan even obtained hybrids between Asterias, a starfish, andArbacia, a sea-urchin, a cross as remote as would be that between a fish and a mammal. Vernon got many hybrids by fertilizing the eggs ofStrongylocentrotus lividuswith the sperm ofSphaerechinus granularis. Standfuss has carried on an enormous series of experiments with Lepidopterous insects, and has obtained a very large series of hybrids, of which he has kept careful record. Lepidopterists generally begin to suspect that many curious forms offered by dealers as new species are products got by crossing known species. Apellö has succeeded with Teleostean fish; Gebhardt and others with Amphibia. Elliot and Suchetet have studied carefully the question of hybridization occurring normally among birds, and have got together a very large body of evidence. Among the cases cited by Elliot the most striking are that of the hybrid betweenColaptes caferandC. auratus, which occurs over a very wide area of North America and is known asC. hybridus, and the hybrid betweenEuplocamus lineatusandE. horsfieldi, which appears to be common in Assam. St M. Podmore has produced successful crosses between the wood-pigeon (Columba palumbus) and a domesticated variety of the rock pigeon (C. livia). Among mammals noteworthy results have been obtained by Professor Cossar Ewart, who has bred nine zebra hybrids by crossing mares of various sizes with a zebra stallion, and who has studied in addition three hybrids out of zebra mares, one sired by a donkey, the others by ponies. Crosses have been made between the common rabbit (Lepus cuniculus) and the guinea-pig (Cavia cobaya), and examples of the results have been exhibited in the Zoological Gardens of Sydney, New South Wales. The Carnivora generally are very easy to hybridize, and many successful experiments have been made with animals in captivity. Karl Hagenbeck of Hamburg has produced crosses between the lion (Felis leo) and the tiger (F. tigris). What was probably a “tri-hybrid” in which lion, leopard and jaguar were mingled was exhibited by a London showman in 1908. Crosses between various species of the smaller cats have been fertile on many occasions. The black bear (Ursus americanus) and the European brown bear (U. arctos) bred in the London Zoological Gardens in 1859, but the three cubs did not reach maturity. Hybrids between the brown bear and the grizzly-bear (U. horribilis) have been produced in Cologne, whilst at Halle since 1874 a series of successful matings of polar (U. maritimus) and brown bears have been made. Examples of these hybrid bears have been exhibited by the London Zoological Society. The London Zoological Society has also successfully mated several species of antelopes, for instance, the water-bucksKobus ellipsiprymnusandK. unctuosus, and Selous’s antelopeLimnotragus selousiwithL. gratus.
Focke, for instance, cites cases where hybrids were obtained betweenBrassicaandRaphanus,GaliumandAsperula,CampanulaandPhyteuma,VerbascumandCelsia. Among animals, new records and new experiments are almost equally numerous. Boveri has crossedEchinus microtuberculatuswithSphaerechinus granularis. Thomas Hunt Morgan even obtained hybrids between Asterias, a starfish, andArbacia, a sea-urchin, a cross as remote as would be that between a fish and a mammal. Vernon got many hybrids by fertilizing the eggs ofStrongylocentrotus lividuswith the sperm ofSphaerechinus granularis. Standfuss has carried on an enormous series of experiments with Lepidopterous insects, and has obtained a very large series of hybrids, of which he has kept careful record. Lepidopterists generally begin to suspect that many curious forms offered by dealers as new species are products got by crossing known species. Apellö has succeeded with Teleostean fish; Gebhardt and others with Amphibia. Elliot and Suchetet have studied carefully the question of hybridization occurring normally among birds, and have got together a very large body of evidence. Among the cases cited by Elliot the most striking are that of the hybrid betweenColaptes caferandC. auratus, which occurs over a very wide area of North America and is known asC. hybridus, and the hybrid betweenEuplocamus lineatusandE. horsfieldi, which appears to be common in Assam. St M. Podmore has produced successful crosses between the wood-pigeon (Columba palumbus) and a domesticated variety of the rock pigeon (C. livia). Among mammals noteworthy results have been obtained by Professor Cossar Ewart, who has bred nine zebra hybrids by crossing mares of various sizes with a zebra stallion, and who has studied in addition three hybrids out of zebra mares, one sired by a donkey, the others by ponies. Crosses have been made between the common rabbit (Lepus cuniculus) and the guinea-pig (Cavia cobaya), and examples of the results have been exhibited in the Zoological Gardens of Sydney, New South Wales. The Carnivora generally are very easy to hybridize, and many successful experiments have been made with animals in captivity. Karl Hagenbeck of Hamburg has produced crosses between the lion (Felis leo) and the tiger (F. tigris). What was probably a “tri-hybrid” in which lion, leopard and jaguar were mingled was exhibited by a London showman in 1908. Crosses between various species of the smaller cats have been fertile on many occasions. The black bear (Ursus americanus) and the European brown bear (U. arctos) bred in the London Zoological Gardens in 1859, but the three cubs did not reach maturity. Hybrids between the brown bear and the grizzly-bear (U. horribilis) have been produced in Cologne, whilst at Halle since 1874 a series of successful matings of polar (U. maritimus) and brown bears have been made. Examples of these hybrid bears have been exhibited by the London Zoological Society. The London Zoological Society has also successfully mated several species of antelopes, for instance, the water-bucksKobus ellipsiprymnusandK. unctuosus, and Selous’s antelopeLimnotragus selousiwithL. gratus.
The causes militating against the production of hybrids have also received considerable attention. Delage, discussing the question, states that there is a general proportion between sexual attraction and zoological affinity, and in many cases hybrids are not naturally produced simply from absence of the stimulus to sexual mating, or because of preferential mating within the species or variety. In addition to differences of habit, temperament, time of maturity, and so forth, gross structural differences may make mating impossible. Thus Escherick contends that among insects the peculiar structure of the genital appendages makes cross-impregnation impossible, and there is reason to believe that the specific peculiarities of the modified sexual palps in male spiders have a similar result.
The difficulties, however, may not exist, or may be overcome by experiment, and frequently it is only careful management that is required to produce crossing. Thus it has been found that when the pollen of one species does not succeed in fertilizing the ovules of another species, yet the reciprocal cross may be successful; that is to say, the pollen of the second species may fertilize the ovules of the first. H. M. Vernon, working with sea-urchins, found that the obtaining of hybrids depended on the relative maturity of the sexual products. The difficulties in crossing apparently may extend to the chemiotaxic processes of the actual sexual cells. Thus when the spermatozoa of an urchin were placed in a drop of seawater containing ripe eggs of an urchin and of a starfish, the former eggs became surrounded by clusters of the male cells, while the latter appeared to exert little attraction for the alien germ-cells. Finally, when the actual impregnation of the egg is possible naturally, or has been secured by artificial means, the development of the hybrid may stop at an early stage. Thus hybrids between the urchin and the starfish, animals belonging to different classes, reached only the stage of the pluteus larva. A. D. Apellö, experimenting with Teleostean fish, found that very often impregnation and segmentation occurred, but that the development broke down immediately afterwards. W. Gebhardt, crossingRana esculentawithR. arvalis, found that the cleavage of the ovum was normal, but that abnormality began with the gastrula, and that development soon stopped. In a very general fashion there appears to be a parallel between the zoological affinity and the extent to which the incomplete development of the hybrid proceeds.
The difficulties, however, may not exist, or may be overcome by experiment, and frequently it is only careful management that is required to produce crossing. Thus it has been found that when the pollen of one species does not succeed in fertilizing the ovules of another species, yet the reciprocal cross may be successful; that is to say, the pollen of the second species may fertilize the ovules of the first. H. M. Vernon, working with sea-urchins, found that the obtaining of hybrids depended on the relative maturity of the sexual products. The difficulties in crossing apparently may extend to the chemiotaxic processes of the actual sexual cells. Thus when the spermatozoa of an urchin were placed in a drop of seawater containing ripe eggs of an urchin and of a starfish, the former eggs became surrounded by clusters of the male cells, while the latter appeared to exert little attraction for the alien germ-cells. Finally, when the actual impregnation of the egg is possible naturally, or has been secured by artificial means, the development of the hybrid may stop at an early stage. Thus hybrids between the urchin and the starfish, animals belonging to different classes, reached only the stage of the pluteus larva. A. D. Apellö, experimenting with Teleostean fish, found that very often impregnation and segmentation occurred, but that the development broke down immediately afterwards. W. Gebhardt, crossingRana esculentawithR. arvalis, found that the cleavage of the ovum was normal, but that abnormality began with the gastrula, and that development soon stopped. In a very general fashion there appears to be a parallel between the zoological affinity and the extent to which the incomplete development of the hybrid proceeds.
As to the sterility of hybridsinter se, or with either of the parent forms, information is still wanted. Delage, summing up the evidence in a general way, states that mongrels are more fertile and stronger than their parents, while hybrids are at least equally hardy but less fertile. While many of the hybrid products of horticulturists are certainly infertile, others appear to be indefinitely fertile.
Focke, it is true, states that the hybrids betweenPrimula auriculaandP. hirsutaare fertile for many generations, but not indefinitely so; but, while this may be true for the particular case, there seems no reason to doubt that many plant hybrids are quite fertile. In the case of animals the evidence is rather against fertility. Standfuss, who has made experiments lasting over many years, and who has dealt with many genera of Lepidoptera, obtained no fertile hybrid females, although he found that hybrid males paired readily and successfully with pure-bred females of the parent races. Elliot,dealing with birds, concluded that no hybrids were fertile with one another beyond the second generation, but thought that they were fertile with members of the parent races. Wallace, on the other hand, cites from Quatrefages the case of hybrids between the mothsBombyx cynthiaandB. arrindia, which were stated to be fertileinter sefor eight generations. He also states that hybrids between the sheep and goat have a limited fertilityinter se. Charles Darwin, however, had evidence that some hybrid pheasants were completely fertile, and he himself interbred the progeny of crosses between the common and Chinese geese, whilst there appears to be no doubt as to the complete fertility of the crosses between many species of ducks, J. L. Bonhote having interbred in various crosses for several generations the mallard (Anas boschas), the Indian spot-bill duck (A. poecilorhyncha), the New Zealand grey duck (A. superciliosa) and the pin-tail (Dafila acuta). Podmore’s pigeon hybrids were fertileinter se, a specimen having been exhibited at the London Zoological Gardens. The hybrids between the brown and polar bears bred at Halle proved to be fertile, both with one of the parent species and with one another.Cornevin and Lesbre state that in 1873 an Arab mule was fertilized in Africa by a stallion, and gave birth to female offspring which she suckled. All three were brought to the Jardin d’Acclimatation in Paris, and there the mule had a second female colt to the same father, and subsequently two male colts in succession to an ass and to a stallion. The female progeny were fertilized, but their offspring were feeble and died at birth. Cossar Ewart gives an account of a recent Indian case in which a female mule gave birth to a male colt. He points out, however, that many mistakes have been made about the breeding of hybrids, and is not altogether inclined to accept this supposed case. Very little has been published with regard to the most important question, as to the actual condition of the sexual organs and cells in hybrids. There does not appear to be gross anatomical defect to account for the infertility of hybrids, but microscopical examination in a large number of cases is wanted. Cossar Ewart, to whom indeed much of the most interesting recent work on hybrids is due, states that in male zebra-hybrids the sexual cells were immature, the tails of the spermatozoa being much shorter than those of the similar cells in stallions and zebras. He adds, however, that the male hybrids he examined were young, and might not have been sexually mature. He examined microscopically the ovary of a female zebra-hybrid and found one large and several small Graafian follicles, in all respects similar to those in a normal mare or female zebra. A careful study of the sexual organs in animal and plant hybrids is very much to be desired, but it may be said that so far as our present knowledge goes there is not to be expected any obvious microscopical cause of the relative infertility of hybrids.
Focke, it is true, states that the hybrids betweenPrimula auriculaandP. hirsutaare fertile for many generations, but not indefinitely so; but, while this may be true for the particular case, there seems no reason to doubt that many plant hybrids are quite fertile. In the case of animals the evidence is rather against fertility. Standfuss, who has made experiments lasting over many years, and who has dealt with many genera of Lepidoptera, obtained no fertile hybrid females, although he found that hybrid males paired readily and successfully with pure-bred females of the parent races. Elliot,dealing with birds, concluded that no hybrids were fertile with one another beyond the second generation, but thought that they were fertile with members of the parent races. Wallace, on the other hand, cites from Quatrefages the case of hybrids between the mothsBombyx cynthiaandB. arrindia, which were stated to be fertileinter sefor eight generations. He also states that hybrids between the sheep and goat have a limited fertilityinter se. Charles Darwin, however, had evidence that some hybrid pheasants were completely fertile, and he himself interbred the progeny of crosses between the common and Chinese geese, whilst there appears to be no doubt as to the complete fertility of the crosses between many species of ducks, J. L. Bonhote having interbred in various crosses for several generations the mallard (Anas boschas), the Indian spot-bill duck (A. poecilorhyncha), the New Zealand grey duck (A. superciliosa) and the pin-tail (Dafila acuta). Podmore’s pigeon hybrids were fertileinter se, a specimen having been exhibited at the London Zoological Gardens. The hybrids between the brown and polar bears bred at Halle proved to be fertile, both with one of the parent species and with one another.
Cornevin and Lesbre state that in 1873 an Arab mule was fertilized in Africa by a stallion, and gave birth to female offspring which she suckled. All three were brought to the Jardin d’Acclimatation in Paris, and there the mule had a second female colt to the same father, and subsequently two male colts in succession to an ass and to a stallion. The female progeny were fertilized, but their offspring were feeble and died at birth. Cossar Ewart gives an account of a recent Indian case in which a female mule gave birth to a male colt. He points out, however, that many mistakes have been made about the breeding of hybrids, and is not altogether inclined to accept this supposed case. Very little has been published with regard to the most important question, as to the actual condition of the sexual organs and cells in hybrids. There does not appear to be gross anatomical defect to account for the infertility of hybrids, but microscopical examination in a large number of cases is wanted. Cossar Ewart, to whom indeed much of the most interesting recent work on hybrids is due, states that in male zebra-hybrids the sexual cells were immature, the tails of the spermatozoa being much shorter than those of the similar cells in stallions and zebras. He adds, however, that the male hybrids he examined were young, and might not have been sexually mature. He examined microscopically the ovary of a female zebra-hybrid and found one large and several small Graafian follicles, in all respects similar to those in a normal mare or female zebra. A careful study of the sexual organs in animal and plant hybrids is very much to be desired, but it may be said that so far as our present knowledge goes there is not to be expected any obvious microscopical cause of the relative infertility of hybrids.
The relative variability of hybrids has received considerable attention from many writers. Horticulturists, as Bateson has written, are “aware of the great and striking variations which occur in so many orders of plants when hybridization is effected.” The phrase has been used “breaking the constitution of a plant” to indicate the effect produced in the offspring of a hybrid union, and the device is frequently used by those who are seeking for novelties to introduce on the market. It may be said generally that hybrids are variable, and that the products of hybrids are still more variable. J. L. Bonhote found extreme variations amongst his hybrid ducks. Y. Delage states that in reciprocal crosses there is always a marked tendency for the offspring to resemble the male parents; he quotes from Huxley that the mule, whose male parent is an ass, is more like the ass, and that the hinny, whose male parent is a horse, is more like the horse. Standfuss found among Lepidoptera that males were produced much more often than females, and that these males paired readily. The freshly hatched larvae closely resembled the larvae of the female parent, but in the course of growth the resemblance to the male increased, the extent of the final approximation to the male depending on the relative phylogenetic age of the two parents, the parent of the older species being prepotent. In reciprocal pairing, he found that the male was able to transmit the characters of the parents in a higher degree. Cossar Ewart, in relation to zebra hybrids, has discussed the matter of resemblance to parents in very great detail, and fuller information must be sought in his writings. He shows that the wild parent is not necessarily prepotent, although many writers have urged that view. He described three hybrids bred out of a zebra mare by different horses, and found in all cases that the resemblance to the male or horse parent was more profound. Similarly, zebra-donkey hybrids out of zebra mares bred in France and in Australia were in characters and disposition far more like the donkey parents. The results which he obtained in the hybrids which he bred from a zebra stallion and different mothers were more variable, but there was rather a balance in favour of zebra disposition and against zebra shape and marking.
“Of the nine zebra-horse hybrids I have bred,” he says, “only two in their make and disposition take decidedly after the wild parent. As explained fully below, all the hybrids differ profoundly in the plan of their markings from the zebra, while in their ground colour they take after their respective dams or the ancestors of their dams far more than after the zebra—the hybrid out of the yellow and white Iceland pony,e.g.instead of being light in colour, as I anticipated, is for the most part of a dark dun colour, with but indistinct stripes. The hoofs, mane and tail of the hybrids are at the most intermediate, but this is perhaps partly owing to reversion towards the ancestors of these respective dams. In their disposition and habits they all undoubtedly agree more with the wild sire.”
“Of the nine zebra-horse hybrids I have bred,” he says, “only two in their make and disposition take decidedly after the wild parent. As explained fully below, all the hybrids differ profoundly in the plan of their markings from the zebra, while in their ground colour they take after their respective dams or the ancestors of their dams far more than after the zebra—the hybrid out of the yellow and white Iceland pony,e.g.instead of being light in colour, as I anticipated, is for the most part of a dark dun colour, with but indistinct stripes. The hoofs, mane and tail of the hybrids are at the most intermediate, but this is perhaps partly owing to reversion towards the ancestors of these respective dams. In their disposition and habits they all undoubtedly agree more with the wild sire.”
Ewart’s experiments and his discussion of them also throw important light on the general relation of hybrids to their parents. He found that the coloration and pattern of his zebra hybrids resembled far more those of the Somali or Grévy’s zebra than those of their sire—a Burchell’s zebra. In a general discussion of the stripings of horses, asses and zebras, he came to the conclusion that the Somali zebra represented the older type, and that therefore his zebra hybrids furnished important evidence of the effect of crossing in producing reversion to ancestral type. The same subject has of course been discussed at length by Darwin, in relation to the cross-breeding of varieties of pigeons; but the modern experimentalists who are following the work of Mendel interpret reversion differently (seeMendelism).
Graft-Hybridism.—It is well known that, when two varieties or allied species are grafted together, each retains its distinctive characters. But to this general, if not universal, rule there are on record several alleged exceptions, in which either the scion is said to have partaken of the qualities of the stock, the stock of the scion, or each to have affected the other. Supposing any of these influences to have been exerted, the resulting product would deserve to be called a graft-hybrid. It is clearly a matter of great interest to ascertain whether such formation of hybrids by grafting is really possible; for, if even one instance of such formation could be unequivocally proved, it would show that sexual and asexual reproduction are essentially identical.
The cases of alleged graft-hybridism are exceedingly few, considering the enormous number of grafts that are made every year by horticulturists, and have been so made for centuries. Of these cases the most celebrated are those of Adam’s laburnum (Cytisus Adami) and the bizzarria orange. Adam’s laburnum is now flourishing in numerous places throughout Europe, all the trees having been raised as cuttings from the original graft, which was made by inserting a bud of the purple laburnum into a stock of the yellow. M. Adam, who made the graft, has left on record that from it there sprang the existing hybrid. There can be no question as to the truly hybrid character of the latter—all the peculiarities of both parent species being often blended in the same raceme, flower or even petal; but until the experiment shall have been successfully repeated there must always remain a strong suspicion that, notwithstanding the assertion and doubtless the belief of M. Adam, the hybrid arose as a cross in the ordinary way of seminal reproduction. Similarly, the bizzarria orange, which is unquestionably a hybrid between the bitter orange and the citron—since it presents the remarkable spectacle of these two different fruits blended into one—is stated by the gardener who first succeeded in producing it to have arisen as a graft-hybrid; but here again a similar doubt, similarly due to the need of corroboration, attaches to the statement. And the same remark applies to the still more wonderful case of the so-called trifacial orange, which blends three distinct kinds of fruit in one, and which is said to have been produced by artificially splitting and uniting the seeds taken from the three distinct species, the fruits of which now occur blended in the triple hybrid.
The other instances of alleged graft-hybridism are too numerous to be here noticed in detail; they refer to jessamine, ash, hazel, vine, hyacinth, potato, beet and rose. Of these the cases of the vine, beet and rose are the strongest as evidence of graft-hybridization, from the fact that some of them were producedas the result of careful experiments made by very competent experimentalists. On the whole, the results of some of these experiments, although so few in number, must be regarded as making out a strong case in favour of the possibility of graft-hybridism. For it must always be remembered that, in experiments of this kind, negative evidence, however great in amount, may be logically dissipated by a single positive result.
Theory of Hybridism.—Charles Darwin was interested in hybridism as an experimental side of biology, but still more from the bearing of the facts on the theory of the origin of species. It is obvious that although hybridism is occasionally possible as an exception to the general infertility of species inter se, the exception is still more minimized when it is remembered that the hybrid progeny usually display some degree of sterility. The main facts of hybridism appear to lend support to the old doctrine that there are placed between all species the barriers of mutual sterility. The argument for the fixity of species appears still stronger when the general infertility of species crossing is contrasted with the general fertility of the crossing of natural and artificial varieties. Darwin himself, and afterwards G. J. Romanes, showed, however, that the theory of natural selection did not require the possibility of the commingling of specific types, and that there was no reason to suppose that the mutation of species should depend upon their mutual crossing. There existed more than enough evidence, and this has been added to since, to show that infertility with other species is no criterion of a species, and that there is no exact parallel between the degree of affinity between forms and their readiness to cross. The problem of hybridism is no more than the explanation of the generally reduced fertility of remoter crosses as compared with the generally increased fertility of crosses between organisms slightly different. Darwin considered and rejected the view that the inter-sterility of species could have been the result of natural selection.
“At one time it appeared to me probable,” he wrote (Origin of Species, 6th ed. p. 247), “as it has to others, that the sterility of first crosses and of hybrids might have been slowly acquired through the natural selection of slightly lessened degrees of fertility, which, like any other variation, spontaneously appeared in certain individuals of one variety when crossed with those of another variety. For it would clearly be advantageous to two varieties or incipient species if they could be kept from blending, on the same principle that, when man is selecting at the same time two varieties, it is necessary that he should keep them separate. In the first place, it may be remarked that species inhabiting distinct regions are often sterile when crossed; now it could clearly have been of no advantage to such separated species to have been rendered mutually sterile and, consequently, this could not have been effected through natural selection; but it may perhaps be argued that, if a species were rendered sterile with some one compatriot, sterility with other species would follow as a necessary contingency. In the second place, it is almost as much opposed to the theory of natural selection as to that of special creation, that in reciprocal crosses the male element of one form should have been rendered utterly impotent on a second form, whilst at the same time the male element of this second form is enabled freely to fertilize the first form; for this peculiar state of the reproductive system could hardly have been advantageous to either species.”
“At one time it appeared to me probable,” he wrote (Origin of Species, 6th ed. p. 247), “as it has to others, that the sterility of first crosses and of hybrids might have been slowly acquired through the natural selection of slightly lessened degrees of fertility, which, like any other variation, spontaneously appeared in certain individuals of one variety when crossed with those of another variety. For it would clearly be advantageous to two varieties or incipient species if they could be kept from blending, on the same principle that, when man is selecting at the same time two varieties, it is necessary that he should keep them separate. In the first place, it may be remarked that species inhabiting distinct regions are often sterile when crossed; now it could clearly have been of no advantage to such separated species to have been rendered mutually sterile and, consequently, this could not have been effected through natural selection; but it may perhaps be argued that, if a species were rendered sterile with some one compatriot, sterility with other species would follow as a necessary contingency. In the second place, it is almost as much opposed to the theory of natural selection as to that of special creation, that in reciprocal crosses the male element of one form should have been rendered utterly impotent on a second form, whilst at the same time the male element of this second form is enabled freely to fertilize the first form; for this peculiar state of the reproductive system could hardly have been advantageous to either species.”
Darwin came to the conclusion that the sterility of crossed species must be due to some principle quite independent of natural selection. In his search for such a principle he brought together much evidence as to the instability of the reproductive system, pointing out in particular how frequently wild animals in captivity fail to breed, whereas some domesticated races have been so modified by confinement as to be fertile together although they are descended from species probably mutually infertile. He was disposed to regard the phenomena of differential sterility as, so to speak, by-products of the process of evolution. G. J. Romanes afterwards developed his theory of physiological selection, in which he supposed that the appearance of differential fertility within a species was the starting-point of new species; certain individuals by becoming fertile onlyinter seproceeded along lines of modification diverging from the lines followed by other members of the species. Physiological selection in fact would operate in the same fashion as geographical isolation; if a portion of a species separated on an island tends to become a new species, so also a portion separated by infertility with the others would tend to form a new species. According to Romanes, therefore, mutual infertility was the starting-point, not the result, of specific modification. Romanes, however, did not associate his interesting theory with a sufficient number of facts, and it has left little mark on the history of the subject. A. R. Wallace, on the other hand, has argued that sterility between incipient species may have been increased by natural selection in the same fashion as other favourable variations are supposed to have been accumulated. He thought that “some slight degree of infertility was a not infrequent accompaniment of the external differences which always arise in a state of nature between varieties and incipient species.”
Weismann concluded, from an examination of a series of plant hybrids, that from the same cross hybrids of different character may be obtained, but that the characters are determined at the moment of fertilization; for he found that all the flowers on the same hybrid plant resembled one another in the minutest details of colour and pattern. Darwin already had pointed to the act of fertilization as the determining point, and it is in this direction that the theory of hybridism has made the greatest advance.
The starting-point of the modern views comes from the experiments and conclusions on plant hybrids made by Gregor Mendel and published in 1865. It is uncertain if Darwin had paid attention to this work; Romanes, writing in the 9th edition of thisEncyclopaedia, cited it without comment. First H. de Vries, then W. Bateson and a series of observers returned to the work of Mendel (seeMendelism), and made it the foundation of much experimental work and still more theory. It is still too soon to decide if the confident predictions of the Mendelians are justified, but it seems clear that a combination of Mendel’s numerical results with Weismann’s (seeHeredity) conception of the particulate character of the germ-plasm, or hereditary material, is at the root of the phenomena of hybridism, and that Darwin was justified in supposing it to lie outside the sphere of natural selection and to be a fundamental fact of living matter.