VARIEGATION OF LEAVES.
BY JAMES HOGG.
At the meeting of the Association of Nurserymen in Chicago, last July, one of our prominent horticulturists described leaf variegation as a disease. Incidentally this brought up the question: Does the graft affect the stock upon which it is inserted?
Much confusion of ideas exists upon this subject, largely due to a loose application of the term disease. Strictly speaking, this term is only applicable to that which shows the health of the plant to be impaired. It should be distinguished from aberrant or abnormal forms, for these are not necessarily indicative of disease. Nobody thinks of saying that red or striped roses are diseased because they are departures in color from the white flower of the type species; or that white, yellow, or striped roses are diseased when the color of the type species is red. Nobody thinks of saying that double flowers are evidences of disease in the plant, or that diminution in the size of leaves or variation in their form is a disease. Why then should it be said that because leaves may become of some other color than green, or become party-colored, therefore they are diseased? If it be said that flowers are not leaves, and that therefore the analogy is not a good one, the reply is, that flowers in all their parts, and fruits also, are only leaves differently developed from the type. This fact is a proven one, and so admitted to be by all botanists and vegetable physiologists of the present day. If it be objected that by becoming double, flowers lose the power of reproducing the variety or species, the answer is, that this loss of power is not necessarily the result of disease, but may arise from various other causes. Because an animal is castrated, it surely will not be claimed that therefore it is diseased. In man and in the higher animals the power of reproduction ceases at certain ages, but it cannot therefore be said that such men or animals are diseased. Neither is a redundancy of parts an unequivocal evidence of disease.
Topknot fowls and ducks are as healthy as those which do not have such appendages, and a Shetland pony is as healthy as a Percheron horse, notwithstanding the difference in their size and weight. Again, color in block or in variegation is not positive evidence of disease in animal life. The white Caucasian is as healthy as the negro, the copper-colored Malay as the red Indian. The horse, ox, and hog run through white and red to black both in solid and party-color, and all are equally healthy; so with the rabbit, dog, cat, and others of our domestic animals. In wild animals, birds, reptiles, fishes, and insects, it is the same, so that mere difference in color or combinations of color are not prima facie evidence of disease.
But some will say this may be true of animal life, but not of plant life. That there is a strong and evident analogy, the one with the other, is now universally admitted by physiologists. Formerly many physiologists considered leaf variegation a disease, because it generally ran in stripes lengthwise of the leaf or in spots. In the former case it was supposed to originate from disease in the leaf cells of the leaf stalk, which, as the cells grow longitudinally, naturally prolonged it to the end of the leaf. But the originating of varieties in which the variegation did not assume this form, with other considerations, has done much to upset this theory. In the variegated leaved snowberry we have the center and border of the leaf green, separated the one from the other by an isolated white or yellow zone. In the zebra-leaved eulalia and the zebra-leaved juncus, from Japan, we have the variegation of the leaf transversely instead of longitudinally, so that according to the old theory we have the anomaly of a healthy portion of the leaf producing an unhealthy portion, and that again a healthy one, and thus alternately along the whole length of the leaf.
When we dissect a leaf in its primal development, we find that its cells contain colorless globules, by botanists called chlorophyl or phyto-color; these undergo changes according as they are acted upon by light, oxygen, or other agents, producing green, yellow, red, and other tints. This chlorophyl only exists in the outer or superficial cells of the parenchyma or cellular tissue of the leaf, and thus differs from starch and other substances produced in the internal cells, from which the light is more or less excluded. It is a fatty or wax-like substance, readily dissolved in alcohol or ether. The primal color of all leaves and flowers is white or a pale yellowish hue, as can readily be seen by cutting open a leaf or flower bud. The seed leaves of the French bean are white when they come out of the earth, but they become green an hour afterward under the influence of bright sunshine. A case is on record where in a certain section, some miles in extent, in this country, about the time of the trees coming into leaf, the sun did not shine for twenty days; the leaves developed to nearly their full size, but were of a pale or whitish color; finally, one forenoon the sun shone out fully, and by the middle of the afternoon the trees were in full summer dress. These facts show that the green color of leaves is due to the action of light. Variegation is sometimes produced independently of the chlorophyl, as in Begonia argyrostigma and Carduus marianus, in which it is produced by a layer of air interposed between the epidermis or outer skin of the leaf and the cells beneath; this gives the leaf a bright, silvery appearance.
To what, then, are we to ascribe leaf variegation? I think that it is entirely due to diminished root power; by this I do not mean that the roots are diseased, but that they are either in an aberrant or abnormal state; but disease cannot be predicated upon either of these states. To explain: everybody knows Spirea callosa to be a strong growing shrub, having umbels of rosy-colored flowers and strong, stout roots; the white flowered variety is quite dwarf, is more leafy and bushy than the species, and has more fibrous and delicate roots than the type; the crisp-leaved variety is still more dwarf, very bushy, and very leafy, and has very fine threadlike roots. This would indicate that the aberrance is in the roots; the two varieties are much more leafy in proportion to their size than the species, so that if the leaves controlled the roots, the latter should have been larger in proportion than those of the species. Again, once when, in the autumn, I was preparing my greenhouse plants for their winter quarters, I cut back a "Lady Plymouth" geranium, which chanced to be set away in a cool and somewhat damp cellar. When discovered the following February and started into growth in the greenhouse it produced nothing but solid green leaves, and never afterward produced a variegated leaf. This I attributed to its having gained greater root power during its long season of rest. By this I mean that the roots had grown and greatly increased in size, although there had not been any leaf growth. That roots under certain circumstances do so is well known. The roots of fir trees have been found alive and growing forty five years after the trunks were felled. The same has occurred in an ash tree after its trunk had been sawn off level with the ground. A root of Ipomea sellowii has been known to keep on growing for twelve years after its top had been destroyed by frost; and in all that time it never made buds or leaves, yet it increased to seven times its original weight. The tuberous roots of some of the Tropæolums will continue to grow and increase in size after the tops have been accidentally broken off; and potatoes buried so deep in the earth that they cannot produce tops will produce a crop of new potatoes.
On the other hand, I have had an oak-leaved geranium overlooked in a corner of the greenhouse until it was almost dried up for lack of water. When its branches were pruned back and it was started into growth only one branch showed the almost black center of the leaf, all the rest were clear green. This was an evident case of diminished root power, but the plant grew as thriftily as ever. The lack of the dark marking in the leaves was equivalent to the variegation in other varieties, only in a reverse direction.