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Charles Robert Darwin (1809–1882).  Origin of Species.
The Harvard Classics.  1909–14.
 
V. Laws of Variation
 
Correlated Variation
 
 
I MEAN by this expression that the whole organisation is so tied together during its growth and development, that when slight variations in any one part occur, and are accumulated through natural selection, other parts become modified. This is a very important subject, most imperfectly understood, and no doubt wholly different classes of facts may be here easily confounded together. We shall presently see that simple inheritance often gives the false appearance of correlation. One of the most obvious real cases is, that variations of structure arising in the young or larvæ naturally tend to affect the structure of the mature animal. The several parts of the body which are homologous, and which, at an early embryonic period, are identical in structure, and which are necessarily exposed to similar conditions, seem eminently liable to vary in a like manner: we see this in the right and left sides of the body varying in the same manner; in the front and hind legs, and even in the jaws and limbs, varying together, for the lower jaw is believed by some anatomists to be homologous with the limbs. These tendencies, I do not doubt, may be mastered more or less completely by natural selection; thus a family of stags once existed with an antler only on one side; and if this had been of any great use to the breed, it might probably have been rendered permanent by selection.  1
  Homologous parts, as has been remarked by some authors, tend to cohere; this is often seen in monstrous plants: and nothing is more common than the union of homologous parts in normal structures, as in the union of the petals into a tube. Hard parts seem to affect the form of adjoining soft parts; it is believed by some authors that with birds the diversity in the shape of the pelvis causes the remarkable diversity in the shape of their kidneys. Others believe that the shape of the pelvis in the human mother influences by pressure the shape of the head of the child. In snakes, according to Schlegel, the form of the body and the manner of swallowing determine the position and form of several of the most important viscera.  2
  The nature of the bond is frequently quite obscure. Isidore Geoffroy St-Hilaire has forcibly remarked that certain malconformations frequently, and that others rarely, co-exist, without our being able assign any reason. What can be more singular than the relation in cats between complete whiteness and blue eyes with deafness, or between the tortoise-shell colour and the female sex; or in pigeons between their feathered feet and skin betwixt the outer toes, or between the presence of more or less down on the young pigeon when first hatched, with the future colour of its plumage; or, again, the relation between the hair and teeth in the naked Turkish dog, though here no doubt homology comes into play? With respect to this latter case of correlation, I think it can hardly be accidental, that the two orders of mammals which are most abnormal in their dermal covering, viz., Cetacea (whales) and Edentata (armadilloes, scaly ant-eaters, &c.,) are likewise on the whole the most abnormal in their teeth; but there are so many exceptions to this rule, as Mr. Mivart has remarked, that it has little value.  3
  I know of no case better adapted to show the importance of the laws of correlation and variation, independently of utility and therefore of natural selection, than that of the difference between the outer and inner flowers in some compositous and timbelliferous plants. Every one is familiar with the difference between the ray and central florets of, for instance, the daisy, and this difference is often accompanied with the partial or complete abortion of the reproductive organs. But in some of these plants, the seeds also differ in shape and sculpture. These differences have sometimes been attributed to the pressure of the involuera on the florets, or to their mutual pressure, and the shape of the seeds in the ray-florets of some Compositæ countenances this idea; but with the Umbelliferæ, it is by no means, as Dr. Hooker informs me, the species with the densest heads which most frequently differ in their inner and outer flowers. It might have been thought that the development of the ray-petals by drawing nourishment from the reproductive organs causes their abortion; but this can hardly be the sole cause, for in some Compositae the seeds of the outer and inner florets differ, without any difference in the corolla. Possibly these several differences may be connected with the different flow of nutriment towards the central and external flowers: we know, at least, that with irregular flowers, those nearest to the axis are most subject to peloria, that is to become abnormally symmetrical. I may add, as an instance of this fact, and as a striking case of correlation, that in many pelargoniums, the two upper petals in the central flower of the truss often lose their patches of darker colour; and when this occurs, the adherent nectary is quite aborted; the central flower thus becoming peloric or regular. When the colour is absent from only one of the two upper petals, the nectary is not quite aborted but is much shortened.  4
  With respect to the development of the corolla, Sprengel’s idea that the ray-florets serve to attract insects, whose agency is highly advantageous or necessary for the fertilisation of these plants, is highly probable; and if so, natural selection may have come into play. But with respect to the seeds, it seems impossible that their differences in shape, which are not always correlated with any difference in the corolla, can be in any way beneficial: yet in the Umbelliferæ these differences are of such apparent importance—the seeds being sometimes orthospermous in the exterior flowers and cœlospermous in the central flowers,—that the elder De Candolle founded his main divisions in the order on such characters. Hence modifications of structure, viewed by systematists as of high value, may be wholly due to the laws of variation and correlation, without being, as far as we can judge, of the slightest service to the species.  5
  We may often falsely attribute to correlated variation structures which are common to whole groups of species, and which in truth are simply due to inheritance; for an ancient progenitor may have acquired through natural selection some one modification in structure, and, after thousands of generations, some other and independent modification; and these two modifications, having been transmitted to a whole group of descendants with diverse habits, would naturally be thought to be in some necessary manner correlated. Some other correlations are apparently due to the manner in which natural selection can alone act. For instance, Alph. de Candolle has remarked that winged seeds are never found in fruits which do not open; I should explain this rule by the impossibility of seeds gradually becoming winged through natural selection, unless the capsules were open; for in this case alone could the seeds, which were a little better adapted to be wafted by the wind, gain an advantage over others less well fitted for wide dispersal.  6
 

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