...Now I want to beg for assistance for the new edition of "Origin." Nageli himself urges that plants offer many morphological differences, which from being of no service cannot have been selected, and which he accounts for by an innate principle of progressive development. (697/1. Nageli's "Enstehung und Begriff der Naturhistorischen Art." An address delivered at the public session of the Royal Academy of Sciences of Munich, March 28th, 1865; published by the Academy. Darwin's copy is the 2nd edition; it bears signs, in the pencilled notes on the margins, of having been read with interest. Much of it was translated for him by a German lady, whose version lies with the original among his pamphlets. At page 27 Nageli writes: "It is remarkable that the useful adaptations which Darwin brings forward in the case of animals, and which may be discovered in numbers among plants, are exclusively of a physiological kind, that they always show the formation or transformation of an organ to a special function. I do not know among plants a morphological modification which can be explained on utilitarian principles." Opposite this passage Darwin has written "a very good objection": but Nageli's sentence seems to us to be of the nature of a truism, for it is clear that any structure whose evolution can be believed to have come about by Natural Selection must have a function, and the case falls into the physiological category. The various meanings given to the term morphological makes another difficulty. Nageli cannot use it in the sense of "structural"—in which sense it is often applied, since that would mean that no plant structures have a utilitarian origin. The essence of morphology (in the better and more precise sense) is descent; thus we say that a pollen-grain is morphologically a microspore. And this very example serves to show the falseness of Nageli's view, since a pollen-grain is an adaptation to aerial as opposed to aquatic fertilisation. In the 5th edition of the "Origin," 1869, page 151, Darwin discusses Nageli's essay, confining himself to the simpler statement that there are many structural characters in plants to which we cannot assign uses. See Volume I., Letter 207.) I find old notes about this difficulty; but I have hitherto slurred it over. Nageli gives as instances the alternate and spiral arrangement of leaves, and the arrangement of the cells in the tissues. Would you not consider as a morphological difference the trimerous, tetramerous, etc., divisions of flowers, the ovules being erect or suspended, their attachment being parietal or placental, and even the shape of the seed when of no service to the plant.
Now, I have thought, and want to show, that such differences follow in some unexplained manner from the growth or development of plants which have passed through a long series of adaptive changes. Anyhow, I want to show that these differences do not support the idea of progressive development. Cassini states that the ovaria on the circumference and centre of Compos. flowers differ in essential characters, and so do the seeds in sculpture. The seeds of Umbelliferae in the same relative positions are coelospermous and orthospermous. There is a case given by Augt. St. Hilaire of an erect and suspended ovule in the same ovarium, but perhaps this hardly bears on the point. The summit flower, in Adoxa and rue differ from the lower flowers. What is the difference in flowers of the rue? how is the ovarium, especially in the rue? As Augt. St. Hilaire insists on the locularity of the ovarium varying on the same plant in some of the Rutaceae, such differences do not speak, as it seems to me, in favour of progressive development. Will you turn the subject in your mind, and tell me any more facts. Difference in structure in flowers in different parts of the same plant seems best to show that they are the result of growth or position or amount of nutriment.
I have got your photograph (697/2. A photograph by Mrs. Cameron.) over my chimneypiece, and like it much; but you look down so sharp on me that I shall never be bold enough to wriggle myself out of any contradiction.
Owen pitches into me and Lyell in grand style in the last chapter of volume 3 of "Anat. of Vertebrates." He is a cool hand. He puts words from me in inverted commas and alters them. (697/3. The passage referred to seems to be in Owen's "Anatomy of Vertebrata," III., pages 798, 799, note. "I deeply regretted, therefore, to see in a 'Historical Sketch' of the Progress of Enquiry into the origin of species, prefixed to the fourth edition of that work (1866), that Mr. Darwin, after affirming inaccurately and without evidence, that I admitted Natural Selection to have done something toward that end, to wit, the 'origin of species,' proceeds to remark: 'It is surprising that this admission should not have been made earlier, as Prof. Owen now believes that he promulgated the theory of Natural Selection in a passage read before the Zoological Society in February, 1850, ("Trans." Volume IV., page 15).'" The first of the two passages quoted by Owen from the fourth edition of the "Origin" runs: "Yet he {Prof. Owen} at the same time admits that Natural Selection MAY {our italics} have done something towards this end." In the sixth edition of the "Origin," page xviii., Darwin, after referring to a correspondence in the "London Review" between the Editor of that Journal and Owen, goes on: "It appeared manifest to the editor, as well as to myself, that Prof. Owen claimed to have promulgated the theory of Natural Selection before I had done so;...but as far as it is possible to understand certain recently published passages (Ibid. {"Anat. of Vert."}, Volume III., page 798), I have either partly or wholly again fallen into error. It is consolatory to me that others find Prof. Owen's controversial writings as difficult to understand and to reconcile with each other, as I do. As far as the mere enunciation of the principle of Natural Selection is concerned, it is quite immaterial whether or no Prof. Owen preceded me, for both of us, as shown in this historical sketch, were long ago preceded by Dr. Wells and Mr. Matthews.")
LETTER 698. TO J.D. HOOKER. Down, December 29th, 1868.
Your letter is quite invaluable, for Nageli's essay (698/1. See preceding Letter.) is so clever that it will, and indeed I know it has produced a great effect; so that I shall devote three or four pages to an answer. I have been particularly struck by your statements about erect and suspended ovules. You have given me heart, and I will fight my battle better than I should otherwise have done. I think I cannot resist throwing the contrivances in orchids into his teeth. You say nothing about the flowers of the rue. (698/2. For Ruta see "Origin," Edition V., page 154.) Ask your colleagues whether they know anything about the structure of the flower and ovarium in the uppermost flower. But don't answer on purpose.
I have gone through my long Index of "Gardeners' Chronicle," which was made solely for my own use, and am greatly disappointed to find, as I fear, hardly anything which will be of use to you. (698/3. For Hooker's projected biological book, see Letter 696.) I send such as I have for the chance of their being of use.
LETTER 699. TO J.D. HOOKER. Down, January 16th {1869}.
Your two notes and remarks are of the utmost value, and I am greatly obliged to you for your criticism on the term. "Morphological" seems quite just, but I do not see how I can avoid using it. I found, after writing to you, in Vaucher about the Rue (699/1. "Plantes d'Europe," Volume I., page 559, 1841.), but from what you say I will speak more cautiously. It is the Spanish Chesnut that varies in divergence. Seeds named Viola nana were sent me from Calcutta by Scott. I must refer to the plants as an "Indian species," for though they have produced hundreds of closed flowers, they have not borne one perfect flower. (699/2. The cleistogamic flowers of Viola are used in the discussion on Nageli's views. See "Origin," Edition V., page 153.) You ask whether I want illustrations "of ovules differing in position in different flowers on the same plant." If you know of such cases, I should certainly much like to hear them. Again you speak of the angle of leaf-divergence varying and the variations being transmitted. Was the latter point put in in a hurry to round the sentence, or do you really know of cases?
Whilst looking for notes on the variability of the divisions of the ovarium, position of the ovules, aestivation, etc., I found remarks written fifteen or twenty years ago, showing that I then supposed that characters which were nearly uniform throughout whole groups must be of high vital importance to the plants themselves; consequently I was greatly puzzled how, with organisms having very different habits of life, this uniformity could have been acquired through Natural Selection. Now, I am much inclined to believe, in accordance with the view given towards the close of my MS., that the near approach to uniformity in such structures depends on their not being of vital importance, and therefore not being acted on by Natural Selection. (699/3. This view is given in the "Origin," Edition VI., page 372.) If you have reflected on this point, what do you think of it? I hope that you approved of the argument deduced from the modifications in the small closed flowers.